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Astrid Schwarz • Kurt Jax
Editors
Ecology Revisited
Reflecting on Concepts, Advancing Science
Ecology Revisited
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Editors
Astrid Schwarz
Institute of Philosophy
Technische Universität Darmstadt
Schloss
64283 Darmstadt
Germany
schwarz@phil.tu-darmstadt.de
Kurt Jax
Department of Conservation Biology
Helmholtz Centre for Environmental
Research (UFZ)
Permoserstr. 15, 04318 Leipzig
Germany
kurt.jax@ufz.de
ISBN 978-90-481-9743-9 e-ISBN 978-90-481-9744-6
DOI 10.1007/978-90-481-9744-6
Springer Dordrecht Heidelberg London New York
Library of Congress Control Number: 2011920689
© Springer Science+Business Media B.V. 2011
No part of this work may be reproduced, stored in a retrieval system, or transmitted in any form or by any
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Printed on acid-free paper
Springer is part of Springer Science+Business Media (www.springer.com)
v
Acknowledgements
This book is the outcome of a plurality of activities and all of them informed the
original idea of an edition and research project “Handbook of Ecological Concepts
(HOEK)”. From the very beginning, we envisioned the HOEK as a collective project
that would provide a forum for exchange and debate of ideas in the rather scattered
field of the history and philosophy of ecology, including theoretical ecology.
Alongside concerns about woolliness, lack of transparency and contradictions in the
use of ecological concepts, another important motivation for creating the HOEK, and
constructing it as a genuine collective project, was our feeling that there was a press-
ing need to investigate the diversity in the history and theory of ecology in Europe,
with its different national traditions, histories and scientific styles. This is true in
particular for the early history of ecology from the late nineteenth century through to
World War II, but pertains also to more recent developments. Such later developments
include, for instance, the different national legislations in biological and environmental
conservation, and the changes initiated by national governmental policies, e.g. mea-
sures for the increased use of renewable energies. Political developments such as
these influence ecological science in the different countries, including the selection
and development of concepts and the specific practices of ecology. Equally, however,
the dynamics of concepts and theories also influence politics. This two-way
interaction occurs, for example, in the context of specific implementation strategies
for pan-European laws and directives, such as the European Water Framework
Directive, as well as in the far-reaching and rapid changes in land use patterns and
large-scale ecological restoration projects which started after the political upheavals
of the 1990s, especially in East Germany and Eastern European countries.
In placing such specific emphasis on the European dimension of ecology and the
framing of environmental issues in general – without, of course, excluding other
geographical regions – we were immediately confronted with a rich diversity of
traditions, scientific habits, science policy practices, and languages. Given the addi-
tional fact that the editors’ own native language is German, we had to deal from the
very start with the difficulties – if not impossibilities – and the challenges of trans-
lation, not to mention the point of transition and the semantic hiatus between natu-
ral and technical languages. All this served to raise our awareness even more of the
issues relating to the history of concepts (Begriffsgeschichte) and its importance for
the entire field of ecology and its context of application.
vi
Acknowledgements
Bearing all these complex problems in mind, the project soon took on the
dimensions of a rather gargantuan and unmanageable enterprise. So the first and
most pressing question was how to make this project workable at all. We decided
to start with a mailing campaign, asking our colleagues and friends from ecology,
philosophy, history, linguistics, geography and sociology to give their comments
and opinions about the project. The feedback was overwhelmingly positive. Not
only did they all welcome the project, but by and large most of them got involved
with it, be it as participants in workshops, as members of the editorial board, as
authors, or as reviewers. Our most sincere thanks go out to them. Without their
continuing support the project would never have become a reality. We greatly hope
that this first volume justifies and merits the confidence they had in us.
The project was strongly encouraged and supported from the start by Ludwig
Trepl and Wolfgang Haber from the Lehrstuhl für Landschaftsökologie, Technische
Universität München in Freising-Weihenstephan. Our discussions with them and
several other people from the department helped to sharpen our ideas.
In the project’s very early stages – at the time of the “proto-HOEK”, as it were –
we studied some of the established and excellent working encyclopaedic projects
in Germany very closely. We are grateful to Albrecht von Massow and also to
Markus Bandur who made us familiar with the work of the HmT, the Handwörterbuch
der musikalischen Terminologie.
The post-doctoral project carried out by one of us (Astrid Schwarz) in Paris at
the Maison des Sciences de l’Homme (MSH) and the Institut d’Histoire et de
Philosophie des Sciences et des Techniques (IHPST) enabled the project to take a
great leap forward. My special thanks go to Pascal Acot and also Jean-Pierre Drouin
for spending their time with me in libraries or nice Parisian coffeehouses and repeat-
edly asking important, sometimes inconvenient, but always helpful questions. Many
thanks go also to Patrick Blandin, Donato Bergandi, Serge Frontier, Catherine
Larrère, Patrick Matagne, Denise Pichod-Viale, and a few other colleagues from the
Muséum Nationale d’Histoire Naturelle who engaged with me in many and inspir-
ing discussion. As an outcome of this warm reception, the first workshop of the
project took place in Paris at the MSH, where it was supported especially by
Hinnerk Bruhns and also Caroline zum Kolk.
Two other workshops followed, one in Leipzig (2004) and one in Darmstadt
(2006) both of them generously supported by the Volkswagen Foundation. They
helped to consolidate and extend the “HOEK community” and provided both fur-
ther encouragement and constructive criticism for our joint endeavour.1
Thanks go also to the colleagues at the institutes where we, the editors, are
based. Colleagues from the Institute for Philosophy in Darmstadt were ready to
accept my (Astrid Schwarz) occasionally intense absorption in the project. Petra
Gehring made available her experience as a local editor of the huge editing project
Historisches Wörterbuch der Philosophie, which lasted about 25 years and was
1See www.hoekweb.net for a description of the workshops and a list of participants.
vii
Acknowledgements
completed in 2005. My many conversations with Alfred Nordmann were not only
inspiring but are also reflected directly in my contributions to this volume.
Many colleagues at the Helmholtz Centre for Environmental Research also
contributed their ideas to the project in the course of discussion. They helped me
(Kurt Jax) in particular to anchor our theoretical ideas in the practice of ecology and
its fields of application. A special word of thanks here goes to Klaus Henle, who
accompanied the project with great sympathy and support, not least by providing
the atmosphere and freedom for me to devote time to what is still a rather unusual
topic for an institution dedicated to ecology and environmental conservation, at
least within the German context. Special thanks go posthumously to the philoso-
pher Heidrun Hesse. She was never afraid to go beyond her disciplinary boundaries
and inspired many scientists to reflect on concepts and ideas in ecology and its
applications. Her critical voice will be missed.
We are grateful to the following members of the Editorial Board for their assis-
tance in preparing the book: Pascal Acot, Paris; Sandra Bell, Durham; Patrick
Blandin, Paris; Alexej Ghilarov, Moscow; John Gowdy, Troy, NY; Volker Grimm,
Leipzig; Wolfgang Haber, Freising; Yrjö Haila, Tampere; Getrude Hirsch-Hadorn,
Zürich; Andrew Jamison, Aalborg; Alan Holland, Lancaster; Chunglin Kwa,
Amsterdam; Thomas Potthast, Tübingen; Peter J. Taylor, Boston; Ludwig Trepl,
Freising; Gerhard Wiegleb, Cottbus.
Many authors contributed to this volume whose first language is not English, but
French, Russian, Finnish, Spanish, Norwegian, Italian or German. These manuscripts
have been revised and in some cases translated in their entirety by Kathleen Cross,
Susan Haak, Patrick Hamm, and Paul Ronning. All four worked with unbelievable
care and patience – many thanks to all of them. Special thanks go to Kathleen
Cross, who not only did most of the translation work, but contributed greatly in
many conversations and e-mail exchanges towards strengthening and clarifying a
number of the articles.
Last but not least we want to thank our former collaborator Christian Haak for
his invaluable work in commenting on and editing so many pages of this volume.
He did a wonderful housekeeping job with the information and data management.
The project website can be visited at http://www.hoekweb.net.
Darmstadt Astrid Schwarz
Leipzig Kurt Jax
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ix
Forwarded Foreword2
The initiators of the Handbook of Ecological Concepts (HOEK) argue that the
HOEK should help in clarifying the relation between concepts in scientific ecology
and the objects that are defined by them.3 They do not attempt to tackle the old
metaphysical problem of the world order, nor do they suppose that it is resolved. In
fact, the question of whether a scientist discovers discontinuities, identities or regu-
larities existing independently from himself that are rooted in reality, and from
which he then extracts laws, or whether a scientist explains the world using a con-
ceptual framework developed by himself or others is not a matter of epistemology:
in practice, scientists do observe regularities in nature. Therefore, at the scale of a
human being, oak trees are always, and in the same manner, different from beech
trees and birch trees, etc. At the same time these regularities are prescribed by using
the complex concept “species”, the definition of which relates to meaning ascribed
to the perceived discontinuities.
One can see that this approach to reality is determined by history; the nature of
observations changes over time, as do the meanings of a concept that give sense to
the observations. Consequently, the decision by those responsible for the HOEK to
focus on the meaning of concepts and of what precisely they relate to, by retracing
the process of their construction and their subsequent uses over time, is an innova-
tion in the lexicon of scientific ecology.4 To my knowledge, most of the scientific
2
This is a slightly changed version of a paper given in French on the occasion of the first workshop
of the project HOEK in Paris in 2002. The workshop took place at the Maison des Sciences de
l’Homme Paris and was entitled “HOEK is going to come true”. The editors are very grateful to
Pascal Acot for his generous encouragement on the ground. He accompanied attentively the first
steps of the HOEK, when Astrid Schwarz was a postdoctoral fellow of the DAAD (German
Academic Exchange Service) in Paris.
3
Astrid Schwarz and Kurt Jax, “Outline of the project”, see www.hoek.tu-darmstadt.de
4
Astrid Schwarz and Kurt Jax propose to present general ecological terms (as e.g. ecosystem or
niche) that have developed in this way. Two decades ago a similar project directed by Jacques
Roger (1920–1990) appeared in France. Its failure was its far too great generality and the early
loss of the scientific person in charge (cf. Cahiers pour l’Histoire du Vocabulaire Scientifique,
CNRS-INALF, 1981–1990). Recently a Dictionnaire d’Histoire et Philosophie des Sciences,
edited by Dominique Lecourt (1999), also “generalist” in nature, has been published by Presses
Universitaires de France.
x
thesauri dealing with this domain juxtapose past definitions and different historical
uses of concepts, without really discussing the significance and importance of
changes in signification. Inversely, the historical approach protects against many
ruinous – and let us concede very French – temptations of normalisation and fixa-
tion of the vocabulary.
The project HOEK will certainly confront obstacles. After all, if it were not a
difficult project, others would already have undertaken the risk to carry it out. Let
us begin with the best known but wrong obstacle: the choice of concepts. Most past
attempts have met with criticism on this front, and sometimes justly: the dictionary
of J. Richard Carpenter,5 for example, first published in 1938, does not contain the
word “ecosystem”; the “Vocabulaire d’écologie” of Daget and Godron6 contains
neither the entry “struggle for existence” nor the entry “eutrophication”, nor the
entry “homoeostasis”. Even the current list7 of the HOEK, one might say, shows
some gaps. It does not include, for example, the word “biosphere” but has the entry
“cosmos”. Despite future revisions the project can expect to face criticism of this
kind of fault, be it justified or not. The first thing for our colleagues to do is deter-
mine which words should be in the HOEK that are not yet included, and which
terms are included but should not be …
Some real difficulties will, nevertheless, remain. First, ecologists work within a
historically as well as geographically specific cultural and environmental frame-
work. Their thinking, and therefore the contents of the concepts they are using is
inevitably marked by factors that are not directly scientific. Secondly, ecology is a
discipline in which, for a century, specialists have tended to think in processes
rather than in terms of objects: they speak, for example, of plant successions rather
than of static vegetation. It thus becomes a complex problem to grasp the meaning
of dynamic concepts. In the following I briefly develop these two points.
The Influence of Non-Scientific Factors
Ramon Margalef, the famous Catalan ecologist, remarked that
All schools of ecology are strongly influenced by a genius loci that goes back to the local
landscape [...] The mosaic-like vegetation of the Mediterranean and Alpine countries,
subjected to millennia of human interference, has assisted at the birth of the plant-sociol-
ogy school of Zürich-Montpellier [...] Scandinavia with a poor flora, has produced ecolo-
gists who count every shoot and sprout [...] And it is only natural that the vast spaces and
smooth transitions of North America and Russia have suggested a dynamic approach in
ecology and the theory of climax.8
Forwarded Foreword
5
Carpenter (1938).
6 Daget P, Michel G, with collaboration from David P, Riso J (1974) Vocabulaire d’ecologie.
Hachette, Paris.
7 I refer here to the list of 26.10.2001.
8
Margalef (1968), p. 26 (emphasis P.A.).
xi
Forwarded Foreword
This is not just a joke about the plant sociology of Uppsala (its promoters have been
accused of artificially multiplying the number of associations by using a method of
multistrata analysis of vegetation) or that of Zürich-Montpellier. It is rather a way
of saying that the conceptual systems in ecology are not universal or at least diffi-
cult to generalize: long ago, when I stayed in French Guyana, one of the problems
occupying some botanists there was whether it was possible to apply the method of
minimum area of Josias Braun-Blanquet (1883–1980) to the rich diversity of the
humid tropical primary forest. The answer was it would have been possible theo-
retically, but would have required a sampling area too large to make the method
applicable in practice.
One of the questions for the collaborators of the HOEK might therefore be
whether particular ecological conditions necessarily shape the development of
the system of concepts from which meaning is being defined, or whether it is
possible that universal significance actually exists. This is a crucial question,
because it can address the issue of scientific styles – German, French, etc. – as well
as that of institutional influences.
During the first decade of the twentieth century Charles Flahault (1852–1935),
a French botanist ranger, failed to unify the vocabulary of plant geography because
of the above mentioned problem. At this time, great disorder dominated the nomen-
clature of ecological units. The terms “association” or “formation” were not clearly
defined and were used very differently depending on the respective researcher.
In 1899 the “VII. Internationale Geographenkongress” in Berlin called for a dia-
logue on the vocabulary of vegetation geography and installed a commission to
work on the question. Charles Flahault was invited by Otto Warburg (1859–1938),
Adolf Engler (1844–1930) and the geographer Oscar Drude (1852–1933), but
Flahault claimed to be “unable to come”, probably because of the French-German
conflict during this time concerning the Alsace and Lorraine.
One year later, in Paris this time, Flahault proposed a project for phytogeographi-
cal nomenclature at the first International Botanical Congress (1900). He had estab-
lished a “Nomenclature of geographical and topographical units” (that went from
“le groupe de regions” (group of regions) to “le station” (station) by passing through
all possible intermediates such as “le domaine” (domain), le district (district), le
sous-district (sub-district), etc.). He then attempted to assign to them a correspond-
ing “série des termes phytogéographiques d’ordre biologique” (series of phytogeo-
graphical terms of biological order). Those terms went from the “type de vegetation”
(vegetation type) to the “forme biologique” (biological form), through “groupes
d’associations” (groups of associations), “associations” (associations), etc.
The congress concluded by proposing a huge consultation to be undertaken
through the press and through correspondence. But nothing arose from the consulta-
tion and in 1905, at the “II. Internationaler botanischer Kongress” in Vienna, a
commission, headed by Charles Flahault and the Swiss Carl Schroeter9 (1855–1939),
was named to put forth proposals at the Congress in Brussels in 1910. A comment
9
Carl Schroeter is the inventor of the word autecology (or autoecology), to denote the ecology of
the single isolated plant, and synecology to denote the ecology of an assemblage of plants.
xii
Forwarded Foreword
of the botanist Jules Pavillard (1868–1961) from Montpellier sheds some light on
the appreciation granted to the results of the phytogeographical commission within
the botanist’s community:
Une grosse déception nous attendait à l’issue du IIIe Congrès International de Botanique
tenu à Bruxelles du 14 au 20 mai 1910. La discussion ouverte devant la section de phyto-
géographie n’ a pu conduire à aucune solution définitive des problèmes essentiels de la
nomenclature.10
Beyond the fact previously mentioned, that it was difficult to come to an agreement
between a phytosociologist from Montpellier and one from Scandinavia, important
cultural influences sterilised the final result. The “English Committee” for example –
speaking with one voice – held the position of the botanist Charles Edward Moss
(1870–1930), who was very influential in Great Britain. Moss was influenced by the
work on succession of the North American Frederic Edward Clements (1874–1945).
Moss presumed the dynamics of vegetation to be the essential reality of groups of
vegetation. He proposed to define an association as a “stage in a successive series”
and a formation as “the totality of all stages of a successive series”!
Clements himself had already succumbed to the obsession of neologism in 1902
when he had reacted to the first propositions from Flahault – those of 1900 – in
publishing an article in the journal edited by Heinrich Gustav Adolf Engler
(1844–1930).11 Flahault had recommended using terms limited to particular regions,
according to him not translatable, such as maquis, garrigue, toundra, llanos, etc.
Clements, having a profound systematic – not to mention a little dogmatic – view,
did not miss this nice opportunity; he accused Flahault of forgetting that these
names denote particular types of principal formations one could also find else-
where, and pointed out that some of the terms proposed, such as “ecological series
of groups of associations” or “type of vegetation”, were much too long. Grasping
the opportunity with both hands, he offered his own system, strictly based on Greek
and Latin, as well as a rule for possible neologisms. As one could expect, this
resulted in linguistically monstrous concepts, sometimes pedantic and certainly, for
the most part, never used to this day! He proposed, for example, ochtophilus instead
of “ripicole” or conophorophilus, to denote a plant occuring in coniferous forests.
His project was not carried out: the phytogeographers already had their habits – but
Clements’ criticisms had been noted. And in 1910, because unanimity would have
been necessary to pass a new nomenclature, Flahault’s initiative failed: there were
14% abstentions or “no” on the final vote.
Certainly the HOEK does not intend any normativity in the sense of the project
just reviewed. But the same or similar reasons that made Flahault’s project fail
could influence the development of the project initiated by Astrid Schwarz and Kurt
Jax; or perhaps just the selection of entries that will be finally retained. I think that
we should keep watch over this together.
10
“A big disappointment was awaiting us at the end of the third International Botanical Congress,
held in Brussels between Mai 14th and 20th in 1910. The discussion in the geobotanical section
did not result in any definite solution of the essential problems in nomenclature.”
11
Clements (1902).
xiii
Forwarded Foreword
The Dynamic Point of View in Ecology:
Difficulties and Fertility
The second topic I would like to address is the difficulty in developing a terminology
of movement. A dynamic vocabulary does not denote a being, but, to take up the
old Aristotelian terminology, “a being and a non-being at once”. When I have a
moving object in mind, I am thinking of something that is no longer and at the same
time of something that I believe it will become. To go from a static point of view
to a dynamic perspective in biology means to think in processes rather than in
objects; this can pose serious problems, which I will attempt to demonstrate by
looking closely at the ecology of plant succession.
While at the beginning of the twentieth century most European ecologists
adopted a static perspective, a “photographic” view of a situation, in the United
States a “cinematic” ecology had developed at the start of 1900s. In 1897, in a
pioneering article, the botanist Conway McMillan showed how the physiognomy of
even plant formation could suggest a progressive dynamic: “[Sphagnum moors or
ponds] may be regarded as such glacial ponds or lakes in process of conversion to
forest [...] and almost every imaginable transition may be found from open lakes
with sandy beach-lines continuous on all sides [...] to solid masses of spruce and
tamarack timber.”12 One can see how the physiognomy of succession eventually
suggests the evolution of the vegetation group: the juxtaposition of states caught in
a certain moment in time reflects the process of transformation of the vegetation.
This connection between the physiognomy of a landscape and its successive
development is also expressed in an earlier text of Henry Chandler Cowles
(1869–1939), father of the theory of plant succession: “In the dune region of lake
Michigan, the normal primary formation is the beach; then, in order, the station-
ary beach dunes, the active or wandering dunes, the arrested or transitional dunes,
and the passive or established dunes. The established dunes pass through several
stages, finally culminating in a deciduous mesophytic forest, the normal climax
type in the lake region [...]”.13
Each time, the same type of explanation of movement is proposed: The condi-
tions under which a certain process proceeds is modified by the development of the
process itself: The pioneering vegetation is necessary to reinforce the dunes in
order to assemble the conditions of development of a higher and denser pioneering
vegetation that will then protect the more important shoots of trees and so on. This
type of reflection was very fruitful in the history of ecology. It could explain, for
example, why the vegetation of North America is three times richer in species than
the vegetation of Occidental Europe. During the last glacial epoch, North American
vegetation could recede slowly to Central America and then come back with the
beginning of the current interglacial epoch, because of the North-South orientation
of the Cordilleras. This was not the case in Europe where the chain of the Pyrenees,
the Mediterranean Sea and the Alps formed a barrier difficult to surmount. It was
12McMillan (1896).
13Cowles (1899), p. 20.
xiv
Forwarded Foreword
the introduction of thinking in terms of successional movements that allowed for
this type of analysis (and many others).
But the adoption of a dynamic perspective is very delicate. Certainly, we know
how to represent, or imagine continuous processes. But scientific work is limited to
decomposing the movement into a succession of distinctive, differing states. We
have seen this with plant succession, but the same would apply to, for example, the
morphogenesis of trees. It is necessary to establish discontinuities – but with which
criteria, acceptable for the greatest possible number of scientists, should one estab-
lish the divisions? Once again one is faced with the question concerning thought
styles and scientific schools.
Moreover, the thinking of Clements, who was one of the most important succes-
sion ecologists in the history of ecology, was criticised because of its rigidity. His
organicist conception of communities and their transformations as well as the real-
ity of the climax as an ultimate state – without any turning back – of the succes-
sional evolution, were also criticised vividly in the epoch between the two World
Wars. This is precisely the reason why the nature of ecological systems is still dis-
cussed, and why the word “ecosystem” is so difficult to define. How, in this case,
should one come to an understanding in conceptual questions?
Under the premise that no unifying effort is envisaged, I see the engagement of the
participants of the HOEK as a testimony, undertaking nothing more than an attempt to
understand the reasons for existing controversies. Some of the difficulties we will meet
are not new. I think, without sinking into paranoia, that we will be accused, as I have
mentioned, of adopting a “normative”, “voluntarist” approach, with selections that are
“arbitrary”, “artificial” or even “chauvinist”. But history also teaches us that difficul-
ties and failures are constitutive for scientific progress. So it was that the failure of
1910 (the attempt of Flahault) resulted immediately in the nearly simultaneous founda-
tion of the schools of plant sociology of Uppsala and of Zürich-Montpellier (and much
could be said about them). These schools continued to dominate the European scien-
tific landscape in the field of nomenclature of vegetation groups till the 1950s. And the
justified criticism of them (originating principally from the Anglo-Saxon world)
played an equally important role in the construction of system and dynamics oriented
ecology, which is practiced today in Europe and all over the world.
The HOEK is obviously much more modest than the great attempts at the begin-
ning of the twentieth century to reflect on vocabularies. Nonetheless, in this field
history teaches us at least one thing: it is better to be criticised than to be unproduc-
tive. It follows that the idea of the HOEK initiators and the HOEK participants, in
spite of the difficulties they will have to surmount, is unlike the practice of Charles
Flahault at the beginning of the last century, declaring unashamedly about the con-
cept of (plant) formation: “I never used this word, because I could not decide which
opinion I should side with and which meaning I should give to it; I simply managed
to get along without it!”14
Pascal Acot
14Flahault (1900), p 443.
xv
Forwarded Foreword
References
Carpenter JR (1938) An ecological glossary. The University of Oklahoma Press, Norman
(Reprinted in 1962. Hafner Publishing Company, New-York/London)
Clements FE (1902) A system of nomenclature for phytogeography. Englers Botanische
Jahrbücher 31:1–20
Cowles HC (1899) The ecological relations of the vegetation on the sand dunes of lake Michigan.
The University Press, Chicago, IL
Daget P, Michel G (ed) (1974) Vocabulaire d’ecologie. Hachette, Paris
Flahault C (1900) Projet de nomenclature phytogéographique. Actes du Congrès International de
Botanique, Paris
Lecourt D (ed) (1999) Dictionnaire d’histoire et philosophie des sciences. Presses Universitaires
de France, Paris
Margalef R (1968) Perspectives in ecological theory. University of Chicago Press, Chicago, IL
McMillan C (1896) On the formation of circular muskeags in Tamarack swamps. Bull Torrey
Botanical Club 23:502–503
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xvii
Contents
Part I Design of the Handbook of Ecological Concepts
1 Why Write a Handbook of Ecological Concepts? ............................... 3
Astrid Schwarz and Kurt Jax
2 Structure of the Handbook..................................................................... 11
Kurt Jax and Astrid Schwarz
3 History of Concepts for Ecology ............................................................ 19
Astrid Schwarz
Part II The Foundations of Ecology: Philosophical
and Historical Perspectives
4 Multifaceted Ecology Between Organicism, Emergentism
and Reductionism .................................................................................... 31
Donato Bergandi
5 The Classical Holism-Reductionism Debate in Ecology ...................... 45
Ludwig Trepl and Annette Voigt
Part III About the Inner Structure of Ecology – Some Theses
6 Conceptualizing the Heterogeneity, Embeddedness,
and Ongoing Restructuring That Make Ecological
Complexity ‘Unruly’ ............................................................................... 87
Peter Taylor
7 A Few Theses Regarding the Inner Structure of Ecology ................... 97
Gerhard Wiegleb
8 Dynamics in the Formation of Ecological Knowledge ......................... 117
Astrid Schwarz
xviii Contents
Part IV Main Phases of the History of the Concept “Ecology”
9 Etymology and Original Sources of the Term “Ecology”.................... 145
Astrid Schwarz and Kurt Jax
10 The Early Period of Word and Concept Formation ............................ 149
Kurt Jax and Astrid Schwarz
11 Competing Terms .................................................................................... 155
Kurt Jax and Astrid Schwarz
12 Stabilizing a Concept .............................................................................. 161
Kurt Jax
13 Formation of Scientific Societies ............................................................ 171
Kurt Jax
14 The Fundamental Subdivisions of Ecology ........................................... 175
Kurt Jax and Astrid Schwarz
Part V “Ecology”, Society and the Systems View in the Twentieth
and Twenty-first Century
15 The Rise of Systems Theory in Ecology ................................................ 183
Annette Voigt
16 Ecology and the Environmental Movement.......................................... 195
Andrew Jamison
17 Ecology and Biodiversity at the Beginning of the Twenty-first
Century: Towards a New Paradigm? .................................................... 205
Patrick Blandin
18 An Ecosystem View into the Twenty-first Century ................................ 215
Wolfgang Haber
Part VI Local Conditions of Early Ecology
19 Early Ecology in the German-Speaking World
Through WWII ....................................................................................... 231
Astrid Schwarz and Kurt Jax
20 The History of Early British and US-American
Ecology to 1950 ........................................................................................ 277
Robert McIntosh
xixContents
21 The French Tradition in Ecology: 1820–1950 ...................................... 287
Patrick Matagne
22 Early History of Ecology in Spain, 1868–1936 ..................................... 307
Santos Casado
23 Plant Community, Plantesamfund ........................................................ 325
Peder Anker
24 Looking at Russian Ecology Through the Biosphere Theory ............. 333
Georgy S. Levit
Part VII Border Zones of Scientific Ecology and Other Fields
25 Geography as Ecology ............................................................................ 351
Gerhard Hard
26 Border Zones of Ecology and the Applied Sciences ............................. 369
Yrjö Haila
27 Border Zones of Ecology and Systems Theory ..................................... 385
Egon Becker and Broder Breckling
28 Economy, Ecology and Sustainability ................................................... 405
John M. Gowdy
Picture Credits ................................................................................................. 413
Glossary ........................................................................................................... 415
Author Biography ........................................................................................... 419
Author Index.................................................................................................... 427
Subject Index ................................................................................................... 437
Part I
Design of the Handbook of Ecological
Concepts
3
Ecology has made considerable progress over the last few decades. Huge amounts
of data have been collected and theories, concepts and practices elaborated, greatly
increasing our understanding of living nature and our own influence on it. At several
points during the twentieth century ecology became the focus of high expectations
that it should help to solve the pressing – and now global – environmental problems
that we face, and indeed these expectations appear still to be growing even today.
So is now the right time to write a Handbook of Ecological Concepts that embraces a
philosophical and historical perspective on ecology and its concepts? Is there not rather
a need to produce more ecological data and models pertinent to the environmental
crisis we are experiencing today, with climate change and other global processes of
change? Can we really learn from previous environmental crises, such as Germany’s
“Waldsterben” in the 1980s,1 or the idea of limits to growth, revived in the US of
the 1960s,2 or the discourse on water pollution in the nineteenth century,3 or again the fear
of wood scarcity (“Holznot”) in the eighteenth century?4 Is it really theories and
concepts that play the main organising and disciplining role in science? And, finally,
even if we agree on this, is the seemingly “old-fashioned” form of a handbook the right
Chapter 1
Why Write a Handbook of Ecological
Concepts?
Astrid Schwarz and Kurt Jax
A. Schwarz (*)
Institute of Philosophy, Technische Universität Darmstadt, Schloss, 64283 Darmstadt, Germany
e-mail: schwarz@phil.tu-darmstadt.de
K. Jax
Department of Conservation Biology, Helmholtz Centre for Environmental Research (UFZ),
Permoserstr. 15, 04318 Leipzig, Germany
e-mail: kurt.jax@ufz.de
1 With regard to the “Sterben of the Waldsterben” see, for instance, the conference in July 2007 at
the University of Freiburg (Germany)“Und ewig sterben die Wälder. Das deutsche Waldsterben in
multidisziplinärer Perspektive”. Organised by the Lehrstuhl für Wirtschafts- und Sozialgeschichte
des Historischen Seminars (Franz-Josef Brüggemeier, Jens Ivo Engels) and the Institut für
Forstökonomie (Gerhard Oesten, Roderich von Detten), both University of Freiburg.
2 Höhler (2005); Schwarz (2004); Anker (2005).
3 Kluge (1986); Luckin (1986).
4 See in particular the work of Sieferle on Austria, but also on Switzerland and the UK (Sieferle
et al. 2008).
A. Schwarz and K. Jax (eds.), Ecology Revisited: Reflecting on Concepts, Advancing Science,
DOI 10.1007/978-90-481-9744-6_1, © Springer Science+Business Media B.V. 2011
4 A. Schwarz and K. Jax
way to go about it – in times of collective Wikipedian knowledge production and a
widely acknowledged unravelling of traditional scientific categories and institutions?
We think the time is exactly right for a Handbook of Ecological Concepts.
There are a number of crucial and indeed pressing reasons for acquiring a deeper
understanding not only of ecological concepts but also of the epistemology and history
of ecology as a whole. Research aimed at solving environmental problems as well as
the communication of this research, be it in interdisciplinary dialogue or between
scientists and users in non-academic fields, requires not only evaluating more and
more new data but also establishing clarity and transparency in relation to the concep-
tual foundations of ecology. Ecologists themselves have frequently and repeatedly
deplored the ambiguity and vagueness of concepts.5 A large number of dictionaries of
ecology have been written and there have even been commissions created to debate
ecological terminology.6 The fact that philosophers and historians of ecology have
been quick to join in this chorus of terminological critics is less surprising.7
However, the issue at hand, including the problem of communication, is not just
a matter of terminology, i.e. of finding the right terms or simply defining concepts.
It goes much deeper than this. In order to disentangle the conceptual knots and
strands in ecology and to infuse ecological concepts with greater power, it is neces-
sary to trace the fluctuations and transformations of concepts and the epistemological
questions related to them. This is especially true of traditionally heterogeneous
fields such as the environmental sciences or ecology.
Thus, our handbook is designed to serve a variety of purposes and interests.
Overall, however, the aim of the HOEK is to contribute towards a better under-
standing of the plurality of meanings and epistemological changes in the field of
ecological and environmental knowledge. We argue that ecology and the environ-
mental sciences are not only driven by instruments or experimental techniques but
are primarily organised around concepts and metaphors. It is precisely the concepts
that embody the different epistemic, normative and cultural regimes or styles of
thinking that are important in the field.
It bears noting, though, that our aim is not to provide “correct” definitions of
ecological concepts or to debunk misunderstandings as misconceptions. On the
contrary, we are interested in tracing back and analysing particularly those misun-
derstandings and displacements that are often productive in the transfer of concepts
from one discipline to another or from everyday language to technical language.
Consequently, we are equally concerned to clarify and sharpen concepts by offering
a systematic and historical survey of their different uses and by highlighting the
ways in which concepts are blurred when they transgress disciplinary borders or
even just community borders in ecology itself. The phenomenon of concepts that
5 Looijen (1998); Mayr (1984); Frazier (1994); Peters (1991); Grimm and Wissel (1997); Jax (2006).
6 Such as the working group for phytogeographic nomenclature established by the III. International
Botanical Congress (Flahault and Schröter 1910) or the Committee on Nomenclature created by
the Ecological Society of America in 1931 (Eggleton 1942).
7 Shrader-Frechette and McCoy (1993), or Greg Cooper in a presentation at ISHPSSB in Exeter
2007; see also Sagoff (2003).
51 Why Write a Handbook of Ecological Concepts?
straddle such borders or even transgress them has recently prompted particular
interest and has been addressed from different theoretical positions. Thus, concepts
are variously described as boundary concepts,8 as being nomadic9 or commuting10
between several fields of knowledge; and concepts can function as destabilising
stabilisers11 or immutable mobiles.12
Ecology and the environmental sciences were confronted from the very beginning
with this kind of co-production of the social and the epistemic and with the hybridi-
sation of concepts, objects and institutions. They often failed adequately to fulfil
the work usually required of disciplinary and epistemological purification. One
might say that ecological knowledge is and was produced in a border zone of insti-
tutional, epistemic and metaphysical divisions.13 Consequently ecology has to deal
with openness and indeterminacy in theory building, practices and knowledge pro-
duction in general.
Occasionally, ecologists themselves refer more or less explicitly to this situation
and raise concerns about the quality of their own concepts, calling for greater self-
reflexivity with respect to the conceptual framework of the discipline.14 The lack of
clarity regarding the basic conceptual foundations of ecology actually impedes the
construction of a strong theoretical framework and – to an even greater extent –
communication in inter- and transdisciplinary discourse. Above all, however, there
is considerable pressure from society to clarify concepts, e.g. in the context of
political environmental decisions or legal frameworks that make use of ecological
concepts and knowledge.15 A clearer and more conscious use of concepts will obvi-
ously contribute towards solving environmental problems and improving ecological
research. Thus, as Steward Pimm was able to demonstrate,16 one reason why the
exhaustive debate on the relation between diversity (or complexity) and stability
during the 1960s and 1970s,17 failed to produce any satisfactory outcome was
because it was dealing with several different concepts of “complexity” and “stability”
and thus with several questions instead of just one. These considerations illustrate
8 Star and Griesemer (1989).
9 Stengers (1997).
10 Schwarz in a paper entitled “Commuting concepts and objects in scientific ecology”, paper
given at the first conference of the European Philosophy of Science Conference in Madrid 2007
(http://www.ucm.es/info/epsa07/misc/EPSA07_BookOfAbstracts.pdf).
11 Kaiser and Mayerhauser (2005): “Destabilising Stabilisers”, paper presented at the conference
Imaging Nanospace, Bielefeld.
12 Latour (1993).
13 Ecological knowledge has been and still is produced in physiology or forest science laboratories,
in applied and theoretical contexts, and with a philosophical background rooted in systems theory
or in complexity theories, in reductionism, holism, emergentism, as well as in other -isms.
14 E.g. Haila and Järvinen (1982); Peters (1991); Pickett et al. (1994/2007).
15 For example, the various Ecosystem Management approaches or the Convention on Biological
Diversity and its “Ecosystem Approach”.
16 Pimm (1984).
17 Goodman (1975); Trepl (1995).
6 A. Schwarz and K. Jax
the point that both internal and external reasons exist that not only justify but
actually necessitate a project such as the HOEK.
What Is Innovative About the HOEK?
The idea of the HOEK has taken a long time to develop. In our own earlier work as
ecologists we often sought what we are now attempting to present, namely a guide
to the basic concepts of ecology that helps us to understand a whole range of issues,
including how they originated, what different expressions they take, why there is so
much confusion around some of them, how they influenced the development of
ecological theory and practice, what the problems are in their application, and how
we can best make use of the underlying ideas to create a theoretical ecological
framework and to help solve environmental problems.
The innovative character of the work presented here is that it facilitates rapid access
to the (sometimes multiple) conceptual content of the terms as well as providing in-
depth information about their philosophical and historical context. The structure and
approach of the Handbook (as explained in more detail in Chaps. 2 and 3) differ sub-
stantially from those of common dictionaries of ecology. The most important differ-
ence in comparison with existing reference works is that the HOEK does not seek to
offer short “technical” definitions for alphabetically ordered concepts. Indeed, there
are no standard definitions such as niche being “the functional position of an organism
in its environment, comprising the habitat in which the organism lives, the periods of
time during which it occurs and is active there, and the resources it obtains there”.18
Instead, every meaning of the concept “niche” that has ever occurred is discussed in
its historical context; changes, trends and fashions are elaborated and linked with
persons, institutions, instruments and theories – in short: the concepts are discussed in
an epistemologically explicit space, elucidating the historical, logical and semantic
processes that link a given concept with its object. The Handbook does not cover each
and every term in ecology but only a limited number that are of major theoretical and
practical relevance (see below). In the glossary at the end of the book those concepts
are listed that are relevant in the volume. Being a trilingual catalogue it gives in the
same time an impression of the richness of terms in different languages.
Who Are the Authors?
The overarching goal of this project is to write a philosophically and historically
informed encyclopedic reference work. In calling it a project, we also want to
emphasise that the HOEK is more than a book; it is also an enterprise – indeed we
18 This is the style of definitions given, for instance, in the Concise Oxford Dictionary of Ecology
(Allaby 1994, p. 269).
71 Why Write a Handbook of Ecological Concepts?
might call it an adventure – aimed at bringing together scholars from fields as varied
as ecology, philosophy, history of science, conservation biology, anthropology,
linguistics and other disciplines to chart the field of ecological knowledge. This
they do both by contributing to this book series in various ways – as authors, referees
or advisors – and also by coming together in workshops and projects related to the
topics of the HOEK whose outcomes feed back into it again. The HOEK thus seeks
to serve as a platform to further the self-reflexivity of ecology and the environmental
sciences and to foster the development of a strong theoretical core for these disci-
plines, a core with a sound philosophical basis.
A further important particularity of the HOEK should be mentioned. The treat-
ment of the different concepts of ecology draws strongly on the European history
of ecology. Although this geographical region was decisive for the creation and
early development of ecology as a science, it has frequently been neglected in the
past,19 not the least on account of language barriers. Many fascinating ideas from
early ecology await rediscovery – ideas which, if reflected on and made more
widely known, can help us to avoid reinventing the ecological wheel time and again
and may even spawn new conceptual developments and aid current debates on
ecological theory and practice.
Who Are the Intended Readers?
The HOEK is aimed at people interested in ecology and in the wider realm of envi-
ronmental research and management, as well as people in the area of environmental
policy making. Thus, the researcher from the discipline of ecology may use it to look
up a classic reference or an ill-remembered meaning, or use the information to help
in a clearer structuring of complex research questions (or even research projects).
Students can use the Handbook to gain clarification of an unfamiliar term or to
improve their understanding of the conceptual foundations of ecology. Specialists
looking from a different perspective (such as policy making, management or law)
will also find information for their purposes, e.g. the lawyer’s clerk fishing for argu-
ments with which to impugn some miscreant in court. Equally, though, the HOEK
seeks to exert a powerful theoretical impact and is thus also written for all those
interested in the philosophy and history of ecology and the environmental sciences.
It can contribute to debates aimed at adequately describing engineering or the
applied and fundamental sciences; the HOEK might also turn out to make an impor-
tant contribution to the debate on models and simulation or else offer new insights
in relation to the field sciences, which have as yet received relatively little epistemo-
logical and cultural attention in contrast to the laboratory sciences.
19 For instance the collection of seminal papers “Foundations of Ecology” compiled by Real and
Brown in 1991.
8 A. Schwarz and K. Jax
These two goals – reaching a relatively wide and non-homogeneous public and
using a methodology not very common in the field under investigation – require a
degree of systematic reflection on how best to proceed with such a project. The fol-
lowing sections will therefore introduce the general structure of the HOEK and the
ideas behind it (Chap. 2), and then move on in a more theoretical part to present
some thoughts about the history of concepts (Begriffsgeschichte), its methodology
and its possibilities and limits in a hybrid scientific field such as ecology (Chap. 3).
As we have shown above, then, it is both timely and necessary to write a
Handbook of Ecological Concepts. The field requires systematic and coordinated
treatment. While “wikis” offer an interesting and important new form of common
knowledge production, they do not guarantee the systematic character provided by
a thoroughly edited handbook. This is exactly what the HOEK seeks to offer and
what is needed to better understand and hopefully also improve the theory and
practise of ecology and the environmental sciences. By the same token, we are
persuaded that this necessary and pressing endeavour can be accomplished meth-
odologically and epistemologically in a proper and productive way by not just
referring to but by building on the history and philosophy of science.
References
Allaby M (ed) (1994) The concise Oxford dictionary of ecology. Oxford University Press, Oxford
Anker P (2005) The ecological colonization of space. Environmental history (http://www.
historycooperative.org/cgi-bin/justtop.cgi?act=justtop&url=http://www.historycooperative.
org/journals/eh/10.2/anker.html) (last accessed 10/11/2010)
Eggleton F (1942) Report of committee on nomenclature. Ecology 23:255–257
Flahault Ch, Schröter C (eds) (1910) Phytogeographische Nomenklatur. III. Int. Bot. Kongress,
Brüssel 1910. Zürcher & Furrer, Zürich
Frazier JG (1994) The pressure of terminological stresses – urgency of robust definitions in ecol-
ogy. Bull Br Ecol Soc 25:206–209
Goodman D (1975) The theory of diversity-stability relationships in ecology. Q Rev Biol 50:
237–266
Grimm V, Wissel C (1997) Babel, or the ecological stability discussions: an inventory and analysis
of terminology and a guide for avoiding confusion. Oecologia 109:323–334
Haila Y, Järvinen O (1982) The role of theoretical concepts in understanding the ecological the-
atre: a case study on island biogeography. In: Saarinen E (ed) Conceptual issues in ecology.
D. Reidel, Dordrecht, pp 261–278
Höhler S (2005) Raumschiff ‘Erde’: Lebensraumphantasien im Umweltzeitalter. In: Schröder I,
Höhler S (eds) Welt-Räume. Geschichte, Geographie und Globalisierung seit 1900. Campus,
Frankfurt a.M, pp 258–281
Jax K (2006) The units of ecology: definitions and application. Q Rev Biol 81:237–258
Kaiser M, Mayerhauser T (2005) Nano-Images as Destabilizing Stabilizers. Paper given at the
conference “Imaging NanoSpace – Bildwelten der Nanoforschung”, Center for interdisciplin-
ary Research Bielefeld
Kluge T (1986) Wassernöte. Alano, Aachen
Latour B (1993) We have never been modern. Harvard University Press, Cambridge
Looijen RC (1998) Holism and reductionism in biology and ecology: the mutual dependence of
higher and lower level research programme. Kluwer, Dordrecht
91 Why Write a Handbook of Ecological Concepts?
Luckin B (1986) Pollution and control: a social history of the Thames in the ninetheenth century.
Hilger, Bristol
Mayr E (1984) Die Entwicklung der biologischen Gedankenwelt. Vielfalt, Evolution und
Vererbung. Springer, Berlin
Peters RH (1991) A critique for ecology. Cambridge University Press, Cambridge
Pickett STA, Kolasa J, Jones CG (1994/2007) Ecological understanding, 2nd edn. Academic, San
Diego, 2007
Pimm SL (1984) The complexity and stability of ecosystems. Nature 307:321–326
Real LA, Brown JH (eds) (1991) Book foundations of ecology: classic papers with commentaries.
University of Chicago Press, Chicago
Sagoff M (2003) The plaza and the pendulum: two concepts of ecological science. Biol Philos
18:529–552
Schwarz AE (2004) Shrinking the ecological footprint with nanotechnoscience? In: Baird D,
Nordmann A, Schummer J (eds) Discovering the nanoscale. IOS Press, Amsterdam,
pp 203–208
Shrader-Frechette KS, McCoy ED (1993) Method in ecology: strategies for conservation.
Cambridge University Press, Cambridge
Sieferle R-P, Krausmann F, Schandl H (2008) Socio-ecological regime transitions in Austria and
the United Kingdom. Ecol Econ 1:187–201
Star SL, Griesemer JR (1989) Institutional ecology, translations and boundary objects: amateurs
and professionals in Berkeley’s museum of-vertebrate-zoology, 1907–39. Soc Stud Sci
19:387–420
Stengers I (1997) Power and invention: situating science. University of Minnesota Press,
Minnesota
Trepl L (1995) Die Diversitäts-Stabilitäts-Diskussion in der Ökologie. Berichte der Akademie für
Naturschutz und Landschaftspflege. Beiheift 12:35–49
11
The Handbook of Ecological Concepts deals with fundamental terms that are or have
been of theoretical relevance in scientific ecology. They are discussed using an
approach that to some extent builds on the methodology of history of concepts.
Approaches using such a methodology were developed during the second half of the
twentieth century in various encyclopaedic projects in the fields of history, politics,
musicology and philosophy, among others (for a more detailed account, see Schwarz,
Chap. 3 this volume). Rather than providing simple definitions and explanations,
these approaches seek to trace and reconstruct the dynamics of concept building and
conceptual transformation. This is exactly what this Handbook aims to do and is also
reflected in the structure of the first volume. The following thoughts are rather pro-
visional but confidently assume that this first volume will be followed by other
volumes that allow to unfold the already existing blueprint entirely.
In general terms the articles follow a common scheme. This allows both for
quick and easy reference as well as for in-depth analysis that includes both historical
and philosophical analysis of the concepts concerned. Generally, the concepts will
not to be arranged in alphabetical order but in so-called conceptual clusters, such
as “ecological units” or “ecological interactions”, enabling entries to be structured
in terms of both form and content (see below). These projected volumes are to trace
scientific discourses by strictly tracking a particular term, such as “niche” or
“organism”. The articles follow the scheme described below and address one key
concept each.1 Other volumes which, like this first one, deal with particular
Chapter 2
Structure of the Handbook
Kurt Jax and Astrid Schwarz
K. Jax
Department of Conservation Biology, Helmholtz Centre for Environmental Research (UFZ),
Permoserstr. 15, 04318 Leipzig, Germany
e-mail: kurt.jax@ufz.de
A. Schwarz (*)
Institute of Philosophy, Technische Universität Darmstadt, Schloss, 64283 Darmstadt, Germany
e-mail: schwarz@phil.tu-darmstadt.de
1 The second volume is planned to deal with “ecological units” and would address in four entries
the concepts “organism”, “population”, “community” and “ecosystem” as a common conceptual
cluster. The concepts that appear in this volume are given in a multilingual glossary at the end of
the book. About two thirds of these concepts are so-called main concepts. A more complete list of
entries is given at http://www.hoekweb.tu-darmstadt.de. It is so far limited to approximately 200.
A. Schwarz and K. Jax (eds.), Ecology Revisited: Reflecting on Concepts, Advancing Science,
DOI 10.1007/978-90-481-9744-6_2, © Springer Science+Business Media B.V. 2011
12 K. Jax and A. Schwarz
epistemic, ontological, and socio-political issues of ecology, and require a more
flexible approach in order to describe their subjects adequately. Notwithstanding
this flexibility, all entries are characterised by the fundamental (and, for the
Handbook, essential) focus on both historical and philosophical (epistemological)
perspectives on ecology and its concepts.
What Is the Structure of Each Entry?
Each entry in the Handbook comprises five sections. The first offers some brief
philological information and a summary of the full entry. The second section con-
stitutes the main bulk of the entry, providing an overview of the historical and
epistemological patterns and features of the term, which are then explained in
greater detail in the third section. Up to this point, the sections for the entries follow
a structure that is more or less familiar from encyclopaedical handbooks, such as
the Handwörterbuch der musikalischen Terminologie (HmT 1972–2006) or the
Historisches Wörterbuch der Philosophie (1971–2007). However, the fourth sec-
tion is rather unusual: it brings an openness and flexibility to the project by provid-
ing the possibility for commenting on and thereby supplementing articles by other
authors. For instance, it allows experts from different fields to add comments relating
to previous sections of the entry, to insert important cross-references that need to
be discussed in a more extended form and, finally, it can be also used to invite
authors to present counter-arguments that may have emerged in the process of
designing the entries or during the peer review process. The fifth and final section
lists the literature references for the entire entry.
In the following, the common HOEK scheme for each article, or entry, is given
in more detail:
1. The top of the article always consists of a literal translation of the concept and of
the different forms existing in other languages (at least in English, French and
German), a description of the etymology including the pre- and extra-ecological
uses of the term, an account of the sources containing the first use of the concept
in ecology; and finally a summary of the whole article.
2. The main section of the discussion of each key concept is divided into several
sub-sections:
(a) Main phases of the history of the concept.
(b) Brief account of epistemological changes and influences.
3. This section provides detailed explanations of the main elements as described
in 2. Related aspects are addressed while remaining close to the core elements
of the concept in question. Cross–references are established to problems that
occur when the concept is used outside ecology, e.g. in environmental protection
or biological conservation.
4. Comments by other authors on particular aspects of an entry.
5. Sources/literature.
132 Structure of the Handbook
The form and content thus meet the demands of primary research and offer a quick
guide to specific concepts. Readers who are interested mainly in acquiring a brief
overview of a concept will find this in section 1 of the articles, while sections 2 to 4
offer greater depth by way of more detail and theoretical complexity. Perspectives
that offer a close-up view of contrasting opinions and issues around each concept
alternate with “long shots” that present a broader outlook on how and when the term
entered ecological discourse and the way in which ecological concepts might be related
to other historical entities or even epochs.2 Our intention is that switching the focus
back and forth between perspectives rooted in different times and spaces will stimu-
late new insights into the general polyvalent character of ecology.
What Are Conceptual Clusters?
The classical way of structuring the Handbook would be to start with the letter “A”
for “abundance” and proceed through the alphabet towards “W” for “water cycle”.
One thing this would lead to, however, is a great deal of repetition in the description
of concepts that are similar to one another; what it would also do, though, is leave
largely untouched a discussion of the links between related concepts. These are the
first and most straightforward reasons why the Handbook is structured by what we
have called “conceptual clusters”. A conceptual cluster3 assembles concepts with
common properties in terms of their epistemology, their meaning and function in
ecology, and the phenomena they describe. Examples of such clusters are “ecological
units”, which brings together concepts such as population, community, formation,
biogeoconesis and ecosystem, and “ecological interactions”, which includes con-
cepts such as predation, competition and mutualism.
Conceptual clusters make it possible to describe and compare different but
related terms and concepts more efficiently and conveniently. They avoid repeated
discussions of the same conceptual problems for each concept and reduce the com-
plexity created by the existence of a multitude of similar concepts. They may even
help to structure ecological theories, of which concepts are the basic building
blocks. Finally, the process of comparing and contrasting different concepts that
make up a conceptual cluster also contributes towards a better understanding of
each particular concept. Biocoenosis and population, for example, can be distin-
guished very clearly in almost every definition; the same goes for ecosystem and
plant association. Nevertheless, common to all these concepts is that they describe
units relevant to ecological research containing (usually) more than a single indi-
vidual organism4 and that they have all been subject to the same questions during
their histories. Such questions include, for example, whether they are delimited by
2 See for more details in Pomata (1998).
3 The term for this idea, which we previously had called “conceptual fields”, arose during discussions
at the first HOEK workshop in Paris in 2002.
4 Very rarely, definitions of ecosystems require at minimum only one organism (e.g. Stöcker 1979).
14 K. Jax and A. Schwarz
topographical or process-based (“functional”) boundaries, whether interactions
between the elements are necessary – and, if so, to what extent – to call them eco-
logical units.5 In addition, some of the concepts are considered – rightly or wrongly – to
be synonymous or have been developed conceptually from each other (e.g. on
the basis of analogies or oppositions). Finally, different ecological units are often
considered to be connected hierarchically with each other, for example in a nested
hierarchy from the population to the biosphere. All these reasons suggest that it
is useful first to address the different ecological units together before any new
differentiation can be made.
Conceptual Clusters and Semantic Fields
The idea of conceptual clusters, as introduced here, has some affinity with the con-
cept of Wortfelder (semantic fields), which has been developed in semantics. A
brief look at the similarities and differences between the concept of semantic fields
and that of conceptual clusters will help to sharpen our understanding of the latter
and the purposes it can serve.
The concept of semantic fields was first developed by German linguist Jost
Trier.6 It was intended to describe areas, or clusters, within natural languages in
which words belong and are related to a common conceptual domain (Sinnbezirk).
The meanings of the different words in a semantic field were considered to be
determined by their mutual relations rather than through isolated analysis of each
word. These basic assumptions of Trier’s theory of semantic fields continue to hold
even today in the different forms in which the approach is used.7 Conceptual clusters
likewise delimit a conceptual sphere but, unlike semantic fields, are focused less on
the semantic aspects of the terms described and – as the name emphasizes – more
on the conceptual aspects and on the phenomena to which the concepts pertain. In
addition, conceptual clusters do not deal with natural language but with a technical
language. We are dealing here with terms and concepts that were either newly
created, such as the neologism “ecology”, or are used in a technical way, such as
the word “niche”. Also, our conceptual clusters include not only terms from one
language, English, but where appropriate also from other languages, in particular
from German and French. The “same” term sometimes has distinctively different
meanings when used in different (“natural”) languages.
Conceptual clusters also share the assumption of the Wortfeld theory that (eco-
logical) terms and concepts assembled within a cluster can be understood better
5 See Jax et al. (1998); Jax (2006).
6 Trier (1931).
7 See especially Gloning (2002). Other components of Trier’s original theory, however, have since
been largely rejected, in particular the assumption that the words in a semantic field should cover
the whole conceptual sphere exhaustively in a mosaic-like manner, and that it is possible to struc-
ture clearly the entire vocabulary of a language into semantic fields.
152 Structure of the Handbook
when they are viewed in the light of their interrelations with each other. In other
words, the purpose of conceptual clusters is to provide a fertile context for indi-
vidual concepts, in which the meanings of the concepts are able to emerge by
means of contrast, juxtaposition and interconnection, that is, by highlighting the
relations between different concepts.
How to Construct and Use Conceptual Clusters
The difficult question regarding the criteria according to which conceptual clusters
should be defined, along with what qualifies words to be included in a particular
cluster, is one that our concept also shares with the concept of semantic fields.
There are, of course, many different ways to divide up the multitude of ecological
concepts based on content. Conceptual clusters as we conceive of them here bring
together different terms whose meanings are closely related. These may be different
words with an identical meaning (synonyms), identical terms with partially over-
lapping meanings, overarching concepts and specific instances of concepts (such as
“association” or “formation” as a special expression of the overarching concept of
“community”). In some cases, we can even observe the same word having devel-
oped significantly different meanings (such as the word “function” in ecology,
which can denote both “process” and “role”). Such semantic processes may occur
within ecology or in the course of a shift between scientific and extra-scientific uses
and vice versa, for instance through metaphorical usage.8
In some cases – indeed, ideally – these concepts can also be ordered in a hierar-
chical manner. Thus a cluster may have the heading “ecological units”,9 this being
the overarching concept within which – according to the structure of Volume 2 –
four different key concepts are subsumed, namely “individual organism”, “popula-
tion”, “community” and “ecosystem”.10 Below this level are further specialised
concepts (including “biocoenosis”, “association” and “formation” in the case of
“community”) which share the generic meaning of one of the key concepts but have
8 Sometimes it is the scientific word that is derived through a change of meaning from ordinary
language (e.g. the terms “niche” or “guild”), and sometimes ordinary language uses ecological
terms in a metaphorical or broader sense (e.g. the “political ecosystem”).
9 “Ecological units” are defined here as all those units that are objects of ecological research. Unit
is to be understood here as an aggregation of objects (particulars), which are chosen and arranged
according to such criteria that they can be characterised as new and interesting objects in their own
right.
10 In the case of ecological units, the four key concepts in fact closely approximate to “basic level
concepts” as described by cognitive psychology (see, e.g. Medin and Smith 1984, p. 124):
a middle level of categorisation around which most knowledge is organised and which is the
preferred level of usage in communication (Löbner 2003, pp. 274 ff.). Like basic level concepts
in natural languages, “community” or “ecosystem” have different and diverse uses, so it comes as
no surprise that different meanings and ambiguities arise when the basic concept is mistaken for
the more specialised one derived from it.
16 K. Jax and A. Schwarz
more specific characteristics not shared by all the definitions of the key concept.
Sometimes such specialised concepts have specific designations and sometimes not
(simply being called “community”, for example, but having a more specialised
meaning).
The concepts dealt with in the HOEK will identify and discuss such differences
below the level of the basic terms, but will generally not address terms such as
“association” separately.11
Our approach to conceptual clusters covers both the synchronic and the dia-
chronic dimension of concepts, i.e. we also include previous meanings of the terms
we discuss. The dynamics of concepts and their most recent manifestations are
always linked in a network of previous and current concepts and can best be under-
stood within such networks.
Our aim in using this analytical and conceptual framework is
1. To understand the dynamics of concepts and the structure of ecological theory, and
2. To find frameworks that are appropriate for rendering concepts operational, in
the knowledge that different expressions of the “same” concept may be adequate
for the many different contexts in which they are used
The internal structure of conceptual clusters is not fixed in any general way.
Initially the cluster is constituted by a selection of concepts gathered together under
a general heading. The structure of the cluster can in principle be analysed in a
variety of ways. Although arranging ecological concepts into conceptual clusters
always implies certain theoretical assumptions about the overall structure of eco-
logical knowledge, the selection of concepts included within a particular cluster is
not a statement about the usefulness of this or that concept or theory. On the con-
trary, a conceptual cluster will almost always bring together concepts derived from
completely opposing theories. These concepts cannot be applied to the same ques-
tions with unambiguous results.12 As a consequence, our analysis often may not
even lead to a completely consistent (re-)ordering of concepts within a conceptual
cluster. The development of conceptual clusters for ecological concepts is thus to
some extent itself a research activity. Conceptual clusters thus also provide fresh
impetus for conceptual research within ecology and for the improvement of a
general theoretical framework for ecology.
11 The method described here creates a kind of hierarchical structure of concepts. This hierarchy
exists only at the conceptual level, of course, and does not imply a physical hierarchy, as, for
example, in commonly postulated “nested hierarchies” of ecological units (population as part of
communities as part of ecosystems). Note that “population” and “ecosystem” are on the same level
of the conceptual hierarchy of ecological units.
12 For example, “ecological units” includes both the “holistic” community concepts of Clements
and Thienemann as well as the “reductionistic” community concepts of Gleason and Ramensky.
172 Structure of the Handbook
References
Gloning T (2002) Ausprägungen der Wortfeldtheorie. In: Cruse A, Franz H, Michael J, Peter Rolf
L (eds) Lexicology. An international handbook on the nature and structure of words and
vocabularies, vol 2. Walter de Gruyter, Berlin, pp 728–737
Jax K (2006) The units of ecology. Definitions and application. Q Rev Biol 81:237–258
Jax K, Jones CG, Pickett STA (1998) The self-identity of ecological units. Oikos 82:253–264
Löbner S (2003) Semantik. Eine Einführung. De Gruyter, Berlin
Medin DL, Smith E (1984) Concepts and concept formation. Annu Rev Psychol 35:113–138
Pomata G (1998) Close-ups and long shots. In: Medick H, Anne-Charlott T (eds) Geschlechter-
geschichte und allgemeine Geschichte. Herausforderungen und Perspektiven. Göttingen,
Wallstein, pp 57–98
Stöcker G (1979) Ökosystem - Begriff und Konzeption. Archiv für Naturschutz und Landschafts-
forschung 19:157–176
Trier J (1931) Der deutsche Wortschatz im Sinnbezirk des Verstandes: Die Geschichte eines
sprachlichen Feldes. Winter, Heidelberg
19
The Handbook of Ecological Concepts is particularly interested in the ways in
which ecological concepts are used. Its main concern is to trace the dynamics and
continuity of concepts, that is, to analyse processes of conceptual transformation as
well as strategies for rendering concepts robust in both current and historical eco-
logical knowledge. Concepts are not discussed in terms of being either false or true
but rather as being more or less appropriate to their intended task. One important
criterion for the adequacy and usefulness of a concept is its functional efficiency
and operational reliability,1 which hinges on the power of the concept to classify,
characterise and differentiate processes or phenomena. The greater this power of
differentiation, the more robust the concept.
In this book we seek to “follow the concepts”, that is, looking at them as they go
about their work as part of a language game – and not when they are on holiday, as
Ludwig Wittgenstein once noted by way of critiquing philosophical interest in the
various definitions and historical meanings of a concept rather than in its actual use.
The meaning of a word lies in its use in practice; meaning lies in the act of expres-
sion – and not behind it.2 After all, it is the context and use of a technical concept
which determine its meaning, be it one in current use or one used in the past.
This emphasis on use in the genealogy of scientific concepts has been described
elsewhere in terms of a series of discontinuities. Such discontinuities – some of
them highly significant – are encountered from time to time when tracing the his-
tory of certain concepts. This has led philosophers such as Gaston Bachelard and
Georges Canguilhem to the insight that “scientific thinking is constantly reshaping
Chapter 3
History of Concepts for Ecology
Astrid Schwarz
A. Schwarz (*)
Institute of Philosophy, Technische Universität Darmstadt, Schloss, 64283 Darmstadt, Germany
e-mail: schwarz@phil.tu-darmstadt.de
1 Lübbe 2000: 36: “…die Funktionstüchtigkeit eines Begriffs für einigermaßen randscharfe
Unterscheidungs- und Zuordnungsleistungen ist ein besonders wichtiges Kriterium für die
Zweckmäßigkeit eines Begriffs.”
2 As Gordon Baker points out, Wittgenstein seeks to establish a different form of representation in
the thinking of his reader: “speaking and thinking are operating with signs, and it is use that gives
life to ‘dead’ signs” (Baker 2001, p. 16).
A. Schwarz and K. Jax (eds.), Ecology Revisited: Reflecting on Concepts, Advancing Science,
DOI 10.1007/978-90-481-9744-6_3, © Springer Science+Business Media B.V. 2011
20 A. Schwarz
its past, because its character is one of constant revolution” (Canguilhem 1979, p. 18).3
Thus to achieve a better and more accurate understanding of science, it is essential
that philosophers and historians look at conceptual discontinuities rather than
asserting false continuities by producing collections of biographies or “tableaus of
doctrines” in the style of a natural history (ibid., p. 17). It is crucial to understand –
and to render comprehensible – “the extent to which concepts, attitudes and methods
that are outdated nowadays were outdated even in their own day and, consequently,
how far the outdated past remains the past of an activity which continues to require
scientific naming” (ibid., p. 27).
Thus it is not primarily the genesis (introduction and definition) of a concept that
is the key indicator of its usefulness in a scientific context; instead, it is its continual
reprocessing – its adaptive malleability – that makes a concept useful and thus robust.
By pointing out discontinuities rather than continuities, philosophical attention is
focused on the need for constant regeneration and the accommodation of scientific
concepts in different conceptual and theoretical environments. Since the retrospective
study of historical meaning cannot contribute substantially towards establishing solid
evidence for the usefulness and appropriateness of a concept in the present day, it is
the current use of concepts that ought to be observed and analysed.
In light of these considerations one may want to ask more probingly what, if
anything, a historiography of concepts can contribute to the study of scientific con-
cepts. Can it make a contribution even despite its limited focus on the genesis of
concepts and even though it generates historical knowledge about the meaning of
these concepts at a given time rather than knowledge about their use? Or does the
historiography of concepts surpass these limits and offer a blueprint for a historiog-
raphy of concepts in science?
To begin, we might say that the historiography of concepts undermines any bias
we hold towards our own present, our “Gegenwartsbefangenheit” (Lübbe 2000,
p. 41). The historiography of concepts can sensitize us to the fact that the proliferation
of publications, contexts and techniques for the use of scientific concepts affords only
a very narrow and restrictive “present” in which a concept has a stable meaning. It
also establishes a critical distance to the dynamics of current scientific development,
along the lines of the dictum, “being aware of the lessons of history … makes one
wary of the effects of fashion and error” (Horder 1998, p. 186). In a similar vein
philosopher and historian of biology Jane Maienschein argues that “… good science
requires an historical perspective […] we make progress in science […] by looking
back as well as staring immediately forward at the cutting edge” (2000, p. 341).
Our chief task could be to use knowledge about the methods and concepts of the
“outdated past” in order to properly acknowledge an adequate vantage point from which
to identify our own potentially “outdated present” – outdated in the sense of an expired
fashion, nothing more dramatic than an error, a regressive theory, a dead concept.
Another important impact of a historiography of concepts is that it elevates atten-
tion not only for the distinction but for the character of relation between the first
3 If not otherwise marked, translation of citations was done by Kathleen Cross.
213 History of Concepts for Ecology
world of reality and the third world of concepts and ideas, to paraphrase Popper’s
three world model. The historiography of concepts allows to escape “naïve circular
reasoning from word towards object and back” (Koselleck 1998, p. 121).
All this can be considered as an initial illustration of the thesis put forward in the
subtitle of this book, namely that to reflect on concepts serves to advance science.
Building Blocks of a Historical-Systematic Handbook
Accordingly, one critical feature of a historical-systematic handbook of scientific
concepts needs to be that it adopts an attentive and thoughtful attitude to the particu-
lar, contingent present that is reflected in the interrelated concepts and theories of a
research programme. At the same time, a previous historical or colloquial meaning
may still be present in a concept’s use and may therefore prove relevant in the forma-
tion of scientific concepts, even if these meanings are not always expressed.4 They
might inform the special features of character of nature (Gernot Böhme) or the hard
core of a research programme (Imre Lakatos), working behind the scenes of the
epistemic operations and ascribing meaning to nature implicitly (see Schwarz,
Chap. 8, this volume). “The great tradition of a balance of nature, going back to antiq-
uity, imputed to nature homogeneity, constancy, or equilibrium and abhorred thoughts
of extinction and randomness. Order and coherence were commonly believed in
Christian tradition to be characteristic of Divine providence. Such ideas die hard” -
notes historian of ecology Robert McIntosh (1991, p. 26). Previous meanings from
either a scientific or a non-scientific context may also influence the formation of sci-
entific concepts. As Koselleck noted, “a new term may be coined which expresses in
language previously non-existent experiences or expectations. It cannot be so new,
however, that it was not already virtually contained in the respective existing language
and that it does not draw its meaning from the linguistic context” (1998, p. 30).
The neologism “ecosystem” is a good example, in that it draws together meanings
that are implicit in other chosen words (Begriffswörter) already in use, such as holo-
coen or biosystem (for a more detailed discussion, see Jax and Schwarz, Chap. 11,
this volume). The concept “niche” is an example of the power of the pre-conceptual
meanings present in debates about the environmental requirements of an organism
(“place” niche), the “role” of the organism in the community, and the functional
notion of niche (Haefner 1980, p. 125) as a hypervolume in N-dimensional space.
The special task of a historical-systematic handbook of ecological concepts is to
identify the basic concepts in the field of ecological knowledge and to do justice to
the epistemic and institutional pluralism in ecology. In order to do so, one needs
4 The differences between spoken and written language and between different public spaces
become relevant here. A scientist might talk about his or her background assumptions (and even
get them published, for example, in an interview), but they will not publish them in a scientific
journal. They might write down their notes in a lab diary or even publish their seminar or lecture
notes on the internet; but they will not make them available without these “brackets of place”.
22 A. Schwarz
to acknowledge fully that ecological knowledge is also produced outside the
scientific discipline of ecology. This is knowledge which may, conversely, acquire
relevance for institutionalised ecology. In order to do justice to this institutional
openness, ecology has more recently been described as a border zone and thus as a
discipline located constitutively at the border between institutional and epistemic
fields, as both laboratory and field science. Ecology – or “Border Biology” as
Robert Kohler (2002) proposes – is a culture of layers and mosaics. According to
Kohler, the tension between laboratory and field sciences doesn’t simply disappear
by the border practices effectively becoming a new disciplinary core. Instead,
object constitution and theory building “on the border” is retained in ecology to the
same extent as its institutional character qua border discipline is upheld, with all
the attendant difficulties regarding the status of field sciences in the topology of the
sciences. The fact that ecology was conceived of as a “bridging science” from the
start and has retained this motif even today (see Schwarz and Jax, Chap. 19, this
volume) might be regarded as an ongoing attempt to give border constellations a
positive turn. These constellations are rooted partially in 19th century natural phi-
losophy and are characterised by epistemic and ontological antagonisms. It is only
since around the 1980s that there has been a broader, reflexive border discourse that
addresses the innovative and creative character – and above all the ubiquity – of
borders and mixes in science and technology.
A second feature of an ecological handbook with a historical-systematic per-
spective is that it can show that, throughout the history of the discipline, ecologists
have developed a high degree of sensitivity and attention to what they feel to be
conceptual flaws and fuzzy elements in their technical language (see Acot,
Forwarded Foreword, this volume). This is certainly one consequence of the
epistemic and institutional difficulties scientists inevitably come across in ecologi-
cal knowledge production. But it also signals an awareness that in ecology, as in
biology more generally, it is principally concepts and not theories that are at the
centre of epistemic strategies.
The special feature of this historical-systematic handbook of ecological con-
cepts is that the concepts are discussed not in alphabetical order but rather accord-
ing to so-called conceptual clusters (see Jax and Schwarz, Chap. 2, this volume).
The particular advantage of this approach is not only that the relationship between
the chosen term (Begriffswort) and the concept (Begriff) is rendered more flexible
but also that the history of the concept and the history of the terminology can be
brought together to forge a systematic reconstruction of a concept’s use in the eco-
logical field. A further advantage is that the aggregation in conceptual clusters facili-
tates a productive crossover between conceptual history and other methods, such as
the history of ideas (Lovejoy 1948), historical semantics (Busse 1987, Busse et al.
2005), the history of metaphors, and the history of discourse (Bödeker 2002).
Our handbook pursues a programme that is at once hedonistic and ascetic. It is
hedonistic in its use of methods and in its appropriation of various systematic
schemes: while the conceptual history perspective is central, it functions above all
as a method that is open to critical negotiation in relation to the history of discourse
and the history of ideas, with consideration also to historical semantics in linguistics
233 History of Concepts for Ecology
and the history of metaphor. Insights and descriptive tools from the history of science
and philosophy are made use of, as are systematic representations from philosophical
anthropology and the philosophy of history. The handbook is ascetic in its selection
of concepts and, in particular, in its mode of presentation of those concepts: just a
few concepts – those that structure the discipline – are discussed in this way (see
Annexe), and they are presented in accordance with a particular schema (see Jax
and Schwarz, Chap. 2, this volume).
Closing and Opening the Debate
Since the foretelling of its demise a few years ago (see, for example, Gumbrecht
2006), conceptual history is now said to be in a “state of transition” (Müller 2005), as
heralded by the title of a recent edition of the German-language journal Archiv für
Begriffsgeschichte dedicated to the discipline. It is placed in a relationship of produc-
tive tension to metaphorology and epistemology, to the history of objects (things) and
the history of discourse; the issue of things in language and things for language is
raised with renewed emphasis, the pros and cons of its methodological exigencies
explored and examined. Conceptual history is effectively undergoing a revision driven
by cultural studies. The idea is that it should thus “serve to overcome the ubiquitous
barriers to communication that exist between the representatives of different disci-
plines and cultures”.5 The interdisciplinary configuration of the objects of conceptual
history (Müller and Schmieder 2008, p. XII f.) is discovered and trialled as “dispositif
history” (Berg 2008, p. 329) and as a medium of reflexivity per se (Mayer 2007).
Regardless of the extent to which the specifics of these methodological correc-
tions and readjustments in perspective may or may not prove convincing, there is
unanimous agreement about the fact that a conceptual history of scientific concepts
needs to be written differently than a history of “basic historical concepts”, or of
“musicological terminology”,6 and even than the history of a “philosophia peren-
nis” as conceived by Joachim Ritter, the originator of the Historisches Wörterbuch
der Philosophie (Historical Dictionary of Philosophy HWPh). A conceptual history
of the sciences raises questions, above all, as to the transformability and robustness
of concepts in the formation of scientific objects and scientific theory building. It
calls for consideration of the possibilities and opportunities afforded by conceptual
history to reflect on discontinuities, as these are associated, for example, with the
terms “epistemic break” or “paradigm change”. And, not least, it raises the question
5 From a report by Ernst Müller and Falko Schmieder following the workshop “Begriffsgeschichte in
den Naturwissenschaften” (Conceptual history in the sciences), held 9–10 February 2007 at the
Centre for Literary and Cultural Research, Berlin (Archiv für Begriffsgeschichte 49 (2007), p. 210).
6 The Handbook of Musicological Terminology (Handwörterbuch der musikalischen Terminologie
HmT) was one of the big German conceptual history projects, founded by Hans-Heinrich
Eggebrecht (1970). For our handbook we mainly adopted the structure of the articles that offer
different levels of information.
24 A. Schwarz
of whether conceptual history can do justice to the intermeshing of scientific
practice, the objects of science and their concepts (Müller und Schmieder 2007,
p. 210) – and, if so, how.
A History for Scientific Concepts Versus a History
for Biological or Ecological Concepts?
Emphasis is often placed on the importance of a conceptual history for biology in
contrast to, say, physics. This is because the former has no language amenable to
formalisation and therefore no purely relational concepts at its disposal. Georg
Toepfer, author of the three-volume series Historisches Wörterbuch der Biologie
(2009 preprint), claims (following Georges Canguilhem) that biology is a “concept-
centered science” (begriffszentrierte Naturwissenschaft) (see below, p. xii). Indeed,
Georges Canguilhem gives a positive turn to the fact that it is not possible to transfer
biological terminology wholesale into a formalised mathematical language; instead, he
elevates this incompleteness to a privileged feature of biology.
This characterisation of indeterminate concepts, often valid only locally and limited
in their scope and systematisation, applies to ecology as well. It appears to be of little
import whether these are everyday words, such as “niche” and “energy”, or neolo-
gisms, which are apparently more easy to control. The property of generality becomes
relevant in a different way here. The language game “ecosystem” provides an impres-
sive demonstration of the kind of semantic fecundity a concept is capable of generating
which at first appears sterile by definition. The ecosystem has since carved a path
through a large number of different discourses and disciplines, ranging from linguistics,
economics and medicine to descriptions of the social role of innovative technologies.
Given this, it is almost self-evident that concepts originating in other spheres of science
and technology should also be appropriated by the ecological community, “landscape”
and “carrying capacity” being just two examples. Ecological concepts can also take the
reverse route: specialist terminology becomes a part of everyday language once more.
“Biotope” and “biodiversity” are nowadays in evidence everywhere, and in an environ-
mentally aware society everybody knows what an ecological niche or an ecosystem is,
and what dying forests and climate change mean. “Useful terms taken from a major
scientific discipline (can) become the master keys to an epoch” (Pörksen 2002, 15): the
scientific concept becomes a metaphor in non-scientific usage.
Concept or Metaphor?
This popularisation could be described as a powerful extension of potential relations
of similarity, which are now no longer determined by the conceptual cluster of a
scientific discipline, but are rather opened up in an uncontrolled way. Concepts lose
their contextualisation as nodal points in conceptual networks in which they “are
related to one another as superordinate or subordinate, contrasting or correlative
253 History of Concepts for Ecology
concepts” (Dutt 2008, 244). There is now a consensus regarding the fact that metaphors
are indispensable in scientific speech and writing and that there is a close connection
between concept building and metaphorical usage. What remains in dispute, though,
is how the distinction between concept and metaphor can be explained. There is a
tendency in cultural studies circles to abandon the hotly contested difference between
concept and metaphor in favour of metaphor. However, the reason frequently given –
that, after all, many concept words are polysemic and therefore vague – seems neither
necessary nor sufficient to abandon determinate conceptual content in favor of meta-
phorical transfer and hence the concept as a conception or even as a concept – the more
so as one might argue that “vagueness as well as non-ambiguity are determined by
context” (Teichert 2008, 100). It turns out that the conceptual demarcation of meta-
phor and concept itself depends to a large extent on background theoretical assump-
tions and, as such, is also context-driven or, one might argue, is also part of a politics
of words. However, it is one of the doyens of the historiography of concepts, Reinhart
Koselleck, who has pointed out that a concept “must be ambiguous”, thereby high-
lighting this feature as one necessary and applicable to concepts, whereas elsewhere
it would be applied only to metaphors.
(A) concept must remain ambiguous to be capable of being a concept. […] Thus a concept
may be clear-cut, but it must be ambiguous. All concepts in which an entire process is semi-
otically concentrated defy definition; only something which has no history can be defined
(Nietzsche). A concept brings together the diverse array of historical experience and a sum-
mation of theoretical and practical factual references in a context which is only given and
can only be truly experienced as such by means of the concept . […] Each concept sets
certain horizons as well as certain limits to possible experience and conceivable theory.7
Clearly, this is more of a hypothesis about the concept as it qualifies to be used in
Koselleck’s dictionary of basic historical concepts (Geschichtliche Grundbegriffe),8
but it is anything but a theoretically satisfying definition of a concept (Knobloch
1992, p. 8) insofar as there is no theory of a historiography of concepts (Teichert
2008, p. 111). This is also the case for the Historical Dictionary of Philosophy
(HWPh) that explicitly abandoned the idea of an integrating theory in favour of a
pragmatic procedure; Müller points out that the “HWPh offers not so much a history
of concepts in the strict sense, but instead documents a history of applications of
philosophical terms” (2004, p. 9). The result is that the invariance of the concepts
and the continuity of philosophical meanings is accentuated.
7 (Ein Begriff […] muss vieldeutig bleiben, um ein Begriff sein zu können. […] Ein Begriff kann
also klar, muß aber vieldeutig sein. Alle Begriffe, in denen sich ein ganzer Prozeß semiotisch
zusammenfaßt, entziehen sich der Definition; definierbar ist nur, das, was keine Geschichte hat
(Nietzsche). Ein Begriff bündelt die Vielfalt geschichtlicher Erfahrung und eine Summe von theo-
retischen und praktischen Sachbezügen in einem Zusammenhang, der als solcher nur durch den
Begriff gegeben ist und wirklich erfahrbar wird. […] Mit jedem Begriff werden bestimmte
Horizonte, aber auch Grenzen möglicher Erfahrung und denkbarer Theorie gesetzt (Koselleck
1995, p. 119 f.).
8 Obviously, Koselleck is not describing the use of concepts in a research field. What he gives here
instead is, as Bernhard F. Scholz points out, “a very adequate description of the manner in which
concepts (and words) circulate in the strand of conversation which serves in the construction and
maintenance the social reality of the lifeword” (1998, p. 89).
26 A. Schwarz
9For a more extended discussion of the difference between a scientific and a literary metaphor and
of the concept of explanation as a metaphorical re-description, see Schwarz 2003, pp. 265 ff.).
The role and impact of a concept is viewed completely differently when it is
looked at from the perspective of logic or philosophy of science (Busse 1987,
p. 49 ff.). The focus is obviously not so much on delineating the scope and inten-
tion of a concept (something that may indeed turn out to be difficult, if not impos-
sible, given concepts such as democracy or liberty). Instead, attention is drawn
towards the logic of a socio-political discourse that gives these concepts their
attractive mode of formation: “Those (basic) concepts are ennobled which point
to, crystallise and attract contradictions” (Knobloch 1992, p. 12). Knobloch even
presumes that it might be worthwhile to talk instead about “functional elements
of a historically distinct speech act” (ibid.) and thus to abandon the language
game “concept” completely in favour of a more pronounced conception of the
objects of a historiography of concepts. This orientation towards the speech act
and the discursive per se might be one way of coping with the complexity of historical
(and current) constellations and of removing the dilemma of designation.
Another means of mitigating the disputed difference between metaphor and
concept might be to consider both of them as explanatory models. In the sentence
cited above, “concept” might be regarded as being construed precisely in the sense
of an explanatory model: “Each concept sets certain horizons as well as certain
limits to possible experience and conceivable theory.” The conceptual model deter-
mines how empirical data, associated concepts and hypotheses are linked and how
they might eventually develop explanatory power. From the metaphor side, the
conception of the so-called interactive metaphor, as discussed by Mary Hesse
(1980), might fulfil a similar function. The crucial point is first of all that metaphors
are not looked at initially as rendering similarities visible but rather in the sense of
creating them. The metaphor here is not the result but rather the cause of a relation
of similarity. One frequent objection to this, however, is that such a metaphor would
preclude anything further being said about the scope of the metaphorical predica-
tion and would become arbitrary: once having been launched as a scientific model,
it can no longer be controlled, meaning that it can no longer be foreseen which of
the associated concepts and ideas will ultimately be relevant and perhaps become
“conceivable theories”. But because a metaphor must be successful as a scientific
model, it must also somehow be related to scientific action. Therefore, the relation
of similarity cannot be completely arbitrary but is merely unpredictable. It is ulti-
mately this property of the unforeseeable extension and modification of associated
concepts and ideas which gives rise to a positive heuristics. It is possible to sum up
in essence what is meant by the distinction between unpredictable and arbitrary
extension in three points, which in turn come close to being a definition of a scien-
tific concept. A successful metaphor is not bold but rather reserved; it is interactive
and works in different contexts, and it can be proven to be inconsistent or even
wrong.9 The usefulness of scientific metaphor as a model can be measured by the
degree of “interpretative resonance along with a simultaneous internal suitability to
the object” (Debatin 1990, p. 805).
273 History of Concepts for Ecology
In the end we might want to conclude that the controversy concerning the
respective role of metaphors and concepts draws attention to their common features
which are most relevant for the method of tracking their use: (1) the close interrela-
tion between concept building and scientific practice, (2) the rule of reflexion given
either by a concept or by a metaphor, and finally, (3) the role of conceptual clusters
in integrating different types of concepts, theories and metaphors.
References
Baker G (2001) Wittgenstein: concepts or conceptions? Harv Rev Philos IX:7–23
Berg G (2008) Die Geschichte der Begriffe als Geschichte des Wissens. Methodische
Überlegungen zum ‘practical turn’ in der historischen Semantik. In: Müller E, Schmieder F
(eds) Begriffsgeschichte der Naturwissenschaften. Zur historischen und kulturellen Dimension
naturwissenschaftlicher Konzepte. Walter de Gruyter, Berlin, pp 327–343
Bödeker HE (ed) (2002) Begriffsgeschichte, Diskursgeschichte, Metapherngeschichte. Wallstein
Verlag, Göttingen
Busse D (1987) Historische Semantik. Analyse eines Programms. Klett-Cotta, Stuttgart
Busse D, Niehr T, Wengeler M (eds) (2005) Brisante Semantik. Neue Konzepte und Forschung-
sergebnisse einer kulturwissenschaftlichen Linguistik. Max Niemeyer Verlag, Tübingen
Canguilhem G (1979) Wissenschaftsgeschichte und Epistemologie. Gesammelte Aufsätze (trans
by Bischof M, Seutter W), Wolf Lepenies (ed). Suhrkamp, Frankfurt/M
Debatin B (1990) Der metaphorische Code der Wissenschaft. Zur Bedeutung der Metapher in der
Erkenntnis- und Theoriebildung. S Eur J Semiotic Stud 2:793–820
Dutt C (2008) Funktionen der Begriffsgeschichte. In: Müller E, Schmieder F (eds)
Begriffsgeschichte der Naturwissenschaften. Zur historischen und kulturellen Dimension
naturwissenschaftlicher Konzepte. Walter de Gruyter, Berlin, pp 241–252
Eggebrecht H-H (1970) Das Handwörterbuch der musikalischen Terminologie. Archiv für
Begriffsgeschichte 14:114–125
Gumbrecht HU (2006) Dimensionen und Grenzen der Begriffsgeschichte. Wilhelm Fink Velag,
München
Haefner JW (1980) Two metaphors of the niche. Synthese 43:123–153
Hesse M (1980) Revolutions and reconstructions in the philosophy of science. Harvester Press,
Brighton
Horder TJ (1998) Why do scientists need to be historians? Q Rev Biol 73:175–187
Knobloch C (1992) Überlegungen zur Theorie der Begriffsgeschichte aus sprach- und kommu-
nikationswissenschaftlicher Sicht. Archiv für Begriffsgeschichte 35:7–24
Kohler R (2002) Labscape and landscape. The University of Chicago Press, Chicago
Koselleck R (1995) Vergangene Zukunft: Zur Semantik geschichtlicher Zeiten (1st edn 1979).
Suhrkamp, Frankfurt/M
Koselleck R (1998) Social history and Begriffsgeschichte. In: Hampsher-Monk I, Tilmanns K, van
Vree F (eds) History of concepts: comparative perspectives. Amsterdam University Press,
Amsterdam, pp 23–36
Lovejoy AO (1948) Essays in the history of ideas. The John Hopkins Press, Baltimore
Lübbe H (2000) Begriffsgeschichte und Begriffsnormierung. In: Scholtz G (ed) Die
Interdisziplinarität der Begriffsgeschichte. Meiner, Hamburg, pp 31–41
Maienschein J (2000) Why study history for science? Biol Philos 15:339–348
Mayer H (2007) Nomadisch unscharf. Vorschläge zur Begriffsgeschichte der Naturwissenschaften.
Frankfurter Allgemeine Zeitung 14.02.2007
28 A. Schwarz
McIntosh RP (1991) Concept and terminology of homogeneity and heterogeneity in ecology. In:
Kolasa J, Pickett STA (eds) Ecological heterogeneity. Springer, New York, pp 24–46
Müller E (2005) Einleitung. Bemerkungen zu einer Begriffsgeschichte aus kulturwissenschaftli-
cher Perspektive. In: Ernst Müller (ed) Begriffsgeschichte im Umbruch. Archiv für
Begriffsgeschichte, Sonderheft Jg. 2004, pp 9–20
Müller E, Schmieder F (2008) Einleitung. In: Müller E, Schmieder F (eds) Begriffsgeschichte der
Naturwissenschaften. Zur historischen und kulturellen Dimension naturwissenschaftlicher
Konzepte. Walter de Gruyter, Berlin, pp 11–23
Müller E, Schmieder F (2007) Begriffsgeschichte in den Naturwissenschaften – die historische
Dimension naturwissenschaftlicher Konzepte. Archiv für Begriffsgeschichte 49:210–214
Pörksen U (2002) Die Umdeutung der Geschichte in Natur. Gegenworte 9:12–17
Scholz BF (1998) Conceptual history in context: reconstructing the terminology of an academic
discipline. In: Hampsher-Monk I, Tilmanns K, van Vree F (eds) History of concepts: compara-
tive perspectives. Amsterdam University Press, Amsterdam, pp 87–102
Schwarz AE (2003) Wasserwüste – Mikrokosmos – Ökosystem. Eine Geschichte der Eroberung
des Wasserraumes. Rombach, Freiburg/Br, pp 273–281
Teichert D (2008) Haben naturwissenschaftliche Begriffe eine Geschichte? Anmerkungen zum
Zusammenhang von Metaphorologie und Begriffsgeschichte bei Hans Blumenberg. In: Müller
E, Schmieder F (eds) Begriffsgeschichte der Naturwissenschaften. Zur historischen und kul-
turellen Dimension naturwissenschaftlicher Konzepte. Walter de Gruyter, Berlin, pp 97–116
Toepfer G (forthcoming). Historisches Wörterbuch der Biologie. Geschichte und Theorie der
biologischen Grundbegriffe, Vol 1 (preprint version June 1, 2009). Verlag J.B. Metzler,
Stuttgart
Part II
The Foundations of Ecology: Philosophical
and Historical Perspectives
31
The classical holism-reductionism debate, which has been of major importance to
the development of ecological theory and methodology, is an epistemological
patchwork. At any moment, there is a risk of it slipping into an incoherent, chaotic
Tower of Babel. Yet philosophy, like the sciences, requires that words and their
correlative concepts be used rigorously and univocally. The prevalent use of every-
day language in the holism-reductionism issue may give a false impression regarding
its underlying clarity and coherence. In reality, the conceptual categories underlying
the debate have yet to be accurately defined and consistently used. There is a need
to map out a clear conceptual, logical and epistemological framework.
To this end, we propose a minimalist epistemological foundation. The issue is
easier to grasp if we keep in mind that holism generally represents the ontological
background of emergentism, but does not necessarily coincide with it. We therefore
speak in very loose terms of the “holism-reductionism” debate, although it would really
be better characterised by the terms emergentism and reductionism. The confrontation
between these antagonistic paradigms unfolds at various semantic and operational
levels. In definitional terms, there is not just emergentism and reductionism, but various
kinds of emergentisms and reductionisms. In fact, Ayala (1974; see also Ruse 1988;
Mayr 1988; Beckermann et al. 1992; Jones 2000) have proposed a now classic trilogy
among various semantic domains – ontology, methodology and epistemology. This
trilogy has been used as a kind of epistemological screen to interpret the reductionist
field. It is just as meaningful and useful, however, to apply the same trilogy to the emer-
gentist field. By revealing the basic assumptions of each, we should be better able to
understand the points that are similar and shared, as well as the incommensurable ones.
The first question regarding the emergentism and reductionism debate concerns
the type of explanation the sciences are seeking. At present in the sciences – from
physics to the human sciences – the ontological and epistemological foundation is
essentially naturalistic and materialistic, meaning that all natural (or social) objects,
events and processes can be understood without reference to extra- or supernatural
Chapter 4
Multifaceted Ecology Between Organicism,
Emergentism and Reductionism
Donato Bergandi
D. Bergandi (*)
Muséum National d’Histoire Naturelle, Paris, France
e-mail: bergandi@mnhn.fr
A. Schwarz and K. Jax (eds.), Ecology Revisited: Reflecting on Concepts, Advancing Science,
DOI 10.1007/978-90-481-9744-6_4, © Springer Science+Business Media B.V. 2011
32 D. Bergandi
(vitalistic or theological) entities, causes, aims or explanations. The order and laws
structuring natural reality are intelligible and, in principle, there is no limit to natu-
ralistic explanations. The existence of this philosophical substrate – the existence
of a scientific and naturalistic epistemology – should be taken into account every
time the key words ‘emergentism’ and ‘reductionism’ appear.
In ecology – and, without exception, in the other natural and human sciences – the
classical confrontation between emergentism and reductionism plays a very impor-
tant and structuring role. It is necessary to be aware that their basic assumptions
involve different and generally antinomian ontologies (worldviews, the “true”
structures of reality, or in other words, our “bets” on the structure of reality), meth-
odologies (research strategies) and epistemologies.1 The existence of these specific
semantic domains should be kept in mind every time we approach this issue.
Holism and Reductionism: An Epistemological Confrontation?
Today’s perspective of reductionist cosmological ontology has its antecedents in
the mechanistic worldview of previous centuries. Gradually, from Leucippus and
Democritus to Dalton and, among others, Bohr, reality has been defined from an
atomistic perspective: reality consists of distinct, discrete, indivisible atoms with a
fixed spatio-temporal amplitude. Unlike reductionism, the holistic ontological
perspective of emergentism is continuistic and relational: reality consists of a
continuum of events and processes that are intrinsically interconnected and interde-
pendent. At first sight both reductionism and emergentism currently share a common
scientific philosophy, namely that all biological phenomena are fundamentally
physico-chemical and that the laws of physics and chemistry are applicable to biological
phenomena. Nevertheless, emergentism holds that the various levels of organisation
(physical, biological and psycho-sociological) are characterised by the acquisition
of new and specific properties (emergent properties). These properties increase the
degree of complexity of a given level compared with the various levels of which it
is composed (hierarchical organisation). For this reason, even if physics and chemistry
are normally applicable to, say, ecological phenomena, each level of organisation
requires appropriate laws and theories that allow for an understanding of the
specific properties of that particular level. By contrast, reductionism denies the
existence of emergent properties or else considers them an epiphenomenon strictly
dependent on the state of our knowledge – what is emergent today will lose its
emergent character tomorrow (Hempel and Oppenheim 1948, pp. 149–151).
These ontological assumptions have, of course, significant consequences in the
methodological and epistemological domains. In the methodological domain, the
two perspectives view the analytical method in a very different way.
1 In this context the word ‘epistemology’ connotes the more limited and specific meaning of the
research domain concerning the relationships among theories and laws that belong to different
organisational levels. In other words, it is characterised by the epistemic challenge of “heteroge-
neous reduction”, or “theoretical reductionism” (Ruse 1988).
334 Multifaceted Ecology Between Organicism, Emergentism and Reductionism
Reductionism considers that at a given level of organisation, analytical study of
constituent parts and their relationships is necessary and sufficient to predict, or at
least explain, all the properties of that level. Fundamentally, reductionism is a “bot-
tom-up” strategy. It takes into account the level at which the events to be explained
occur (ecological phenomena, for instance) as well as the lower levels that contribute
to that explanation (for example, genetics, chemistry or physics). An analytical and
additive method, therefore, dissects the entity, or decompose the process, under
examination into its component parts, or phases, and attempts to take into consider-
ation the relationships among them. A successive summation of the individual com-
ponent properties or interactional properties should allow extrapolation of the global
properties of the entity as a whole. In some cases, this dissective and synthetic pro-
cess should allow us to formulate some more general theories or laws.
Methodologically, the emergentist approach, while recognising the need for
analysis, considers its explanatory power limited. In fact, according to an emergen-
tist and hierarchical perspective, the feedback loops that link different levels of
organisation play a role of utmost importance in the determination and causation of
the emergent properties. From a methodological point of view, the higher and lower
levels adjacent to the primary object of study are considered differently than in
methodological reductionism. This approach does not limit the analysis to the con-
stitutive parts of – or their relationships in – a specific level of organisation. In other
words, for this “top-down” approach, both the higher levels (downward causation)
and the lower ones participate in determining the properties of specific levels. Thus,
a multi-level triadic approach – where at least three levels of organisation are con-
sidered simultaneously – is held to be a methodological necessity and is the main
characteristic of the emergentist methodology (Feibleman 1954; Campbell 1974;
Salthe 1985; Bergandi 1995; El-Hani and Pereira 2000).
Epistemologically, reductionism is a mono-directional bottom-up explanatory
strategy. This approach is directly descended from nineteenth century positivism and
from neo-positivism (1920s and 1930s). In its struggle against the intrusiveness of
metaphysics in science, neo-positivism sought a unification of science based on the
language, laws and theories of physics. Epistemological reductionism maintains that
the theories and laws of a specific organisational level can be – and sometimes must
be – “reduced” to the theories and laws of a more “fundamental” field of science
(Woodger 1952; Nagel 1961; Levins and Lewontin 1980; Bunge 1991; Jones 2000).
According to this epistemological perspective, an ideal scientific development will
involve, in the long run, the “de-substantialisation” of non-fundamental sciences. For
instance, taking into account the relationships between ecology (secondary science)
and physics (primary science), ecological laws and theories could be reduced to
physical laws and theories (heterogeneous reduction). Were this to occur, the process
of integration, incorporation and absorption of ecological phenomena in the physical
domain would provide a larger and clearer understanding of all the phenomena that
previously constituted the objects of ecological research. Such a hypothetical reduc-
tion would determine the birth of a new and more meaningful physical science,
emerging from the “dilution” of biology into physics. And, as Popper pointed out,
such a successful reduction is substantially unattainable because it would imply a
“complete” theoretical understanding of life in physical terms (1972).
34 D. Bergandi
Epistemological holism, on the other hand, posits a more dialectical relationship
between laws and theories belonging to different organisational levels. On the one
hand, this perspective holds that there is no scientific domain to which the other
sciences should be reduced. According to emergentist ontology, every organisa-
tional level has one or more emergent properties that are correlated to specific laws
and theories which, in turn, are assumed to be intrinsically non-reducible. On the
other hand, according to Quine (1961, p. 42) “the unit of empirical significance is
the whole of science”.2 This means that the existence of anomalies that cannot be
explained in terms of existing knowledge requires us to make adjustments to science
as a whole. In other words, a transformation in any scientific domain, and not only
in the “fundamental” sciences, can determine changes in any other domain of sci-
ence. This perspective entails rejecting the physical explanation as the fundamental
and preferred form of explanation to which the other sciences have to be reduced.
In sum, it is possible to identify the foundational, philosophical core of all mate-
rialistic emergentist views of reality using the following criteria, which correspond
to different semantic domains:
Ontology
1. Holism: Not all holistic positions are emergentist, but all emergentist views are
holistic. Holism fundamentally means the intrinsic, structural, spatio-temporal
interdependence of phenomena3 and constitutes the major and inescapable onto-
logical presupposition of emergence.
2. Levels of organisation: Reality is a hierarchical, multi-layered, multi-level pro-
cess. According to this interpretation of reality every level of organisation (or
integration) is characterised by specific emergent properties, qualities or behav-
iours. This ontological perspective can be interpreted according to a realistic
view – the levels with their emergent properties definitely represent reality – or
a constructivist one – the levels of organisation are “levels of description” of
reality: we identify levels, and attribute specific properties to them, according to
the purpose of our research.
3. Novelty: The emergent properties of every level of organisation express new
qualities and a new order of phenomena compared with the level of organisation
on which they depend and from which they emerge.
2 It is worth pointing out that, unlike the Quine thesis, the holistic reference of the Duhem thesis
is the whole of physics. Its working has been described according to an organicist perspective: in
physics, as in an organism, all the theories work together, even if they are not all called into play
at the same level of intervention (1977, pp. 187–188).
3 To avoid any risk of misunderstanding, it would be more appropriate to use the term ‘holism’ to
indicate specifically the relational view of reality according to which natural (or social) reality
is constituted by spatio-temporal interdependent entities. Its logical opposite is the ontological
atomistic view.
354 Multifaceted Ecology Between Organicism, Emergentism and Reductionism
Methodology
4. Avoiding the fallacy of “misplaced concreteness” (Whitehead 1925; Dewey and
Bentley 1949). This is a basic prerequisite for any emergentist constructivist meth-
odology. There is a preliminary heuristic assumption that all analytical distinctions
concerning “wholes”, “parts”, and “relations” are pure theoretical “mind construc-
tions” which have meaning only in relationship to the specific aims of the inquiry.
Consequently, wholes, parts and relations must not necessarily be considered to have
an intrinsic ontological reality, merely an epistemic one (see Bergandi 2007).4
5. Multi-level approach: To explain the emergent properties of a specific level of organ-
isation or system, the adjacent lower and higher levels must be considered as signifi-
cant as, and simultaneously with the level of the primary object of research. This
triadic approach is not a luxury but a necessity for any research claiming an emergen-
tist approach. In fact, to restrict to take into consideration the lower level relation-
ships among elements is equivalent to enacting a reductionist methodology.
6. Fallacious attribution of emergent properties: The constructivist background
(see (4) above) should always be borne in mind in the attribution of emergent
properties to a level of organisation. The hypothesis that these properties cannot,
in reality, be effectively attributed to the constituent parts, sub-systems or higher
inclusive levels must be carefully refuted. In fact, any potentially erroneous attri-
bution of an emergent property could result from an incomplete or wrongheaded
analysis of the whole hierarchical structure.
Epistemology
7. Unpredictability: The emergent properties of a level of organisation cannot be
predicted, even in principle, by even the most complete knowledge of the parts,
properties and relationships among the parts.5 In other words, a specific organi-
sation of matter is correlated to exclusive properties. To be able to explain them
would require the constitution of a new or reorganised scientific discipline which
would use new postulates, theories and laws that introduce new terms and pat-
terns suited to the emergent phenomena and properties.6
4 It is worth recalling that constructivism does not deny the existence of a reality (natural, social,
and so on). Rather, this perspective foregrounds the idea that within this reality, thanks to our
epistemic constructs, we identify or recognise certain characteristics, aspects and processes that
are functional to our aims and objectives (scientific, social, and so forth).
5 This is an elliptic formulation; the correct one is the following: the laws concerning the emergent
properties of a level of organisation cannot be predicted, even in principle, by the laws concerning
the lower level relations between the constituent parts.
6 For instance, even the most radical reductionist could not explain biological evolution by refer-
ring only to the overall theoretical package of physics and chemistry; according to Williams: “at least
the one additional postulate of natural selection and its consequence, adaptation, are needed”
(Williams 1966, p. 5; see also 1985, p. 1).
36 D. Bergandi
From Organicism to the Oxymoronic “Reductionist Holism”
of Ecosystem Ecology
From its beginning, ecology has been structured within a holistic ontological frame-
work. Ecology is most widely known as the science that concerns the relationships
between organisms and their environment, that is, a science interested in all the
conditions that permit organisms to live (Haeckel 1866). Early on, this holistic
framework mainly took the form of an organicist worldview. Representative in
this regard are the works of Stephan A. Forbes, Frederic E. Clements and John
Phillips.
Some years after the far-sighted definition of ecology by Ernst Haeckel, Stephan
Alfred Forbes wrote two papers that vividly portrayed the complex, intricate rela-
tionships between organisms and their environments. In On some interactions of
organisms (1880) and The lake as a microcosm (1887; for the concept of “micro-
cosm” as a central metaphor in ecology, see Schwarz 2003), Forbes was among the
first to put forward the idea that natural systems exist in a state of equilibrium and
must be studied “as a whole”. He also delineated a strict connection between natural
selection and the laws of oscillation of plant and animal species. He suggested that
the functional relations among organisms were comparable to the relations between
organs within an animal’s body. Any change (in numbers, habits or distribution)
within a specific plant or animal group will impact various other groups “in a far
extending circle” (1880, p. 3). In the struggle for existence under the influence of
natural selection, predator and prey species ordinarily find a balance and, to a certain
extent, adjust their rates of reproduction accordingly. They have common interests:
an excessive increase in a predator species will inevitably determine a decrease in
the very species that constitute its food supply and consequently a decrease in its own
species. However, Forbes also thought that in the intricate network of relationships
between organisms on the one hand and between organisms and their environments
on the other, the real limits to excessive multiplication of a species are to be found
in the inorganic features of its environment (Ivi, 11, p. 16).
The “lake” was presented by Forbes as the paradigmatic case of a relatively
isolated system in which the “organic complex”, the species assemblage, could not
be studied without taking into account all the forms of relationship between different
species (predator/prey, competition, mutualism, and so forth) belonging to the lake
and the surrounding terrestrial system (1887, p. 537). In other words, prior to the
trophic ecology of Elton (1927) and Lindeman (1942), Forbes considered that when
studying a carnivorous lake fish, one must also take into account the species upon
which it depends for its existence, the organic and inorganic conditions upon which
these species depend, the other competitor species, as well as the entire system of
conditions affecting the existence of the plant and animal species that contribute to
the existence of a specific group of related species (see also Forbes 1914).
The work of other ecologists, including Frederic E. Clements, John Phillips,
Henry A. Gleason and Arthur G. Tansley, shows traces of the influence of a specific
version of what we nowadays call the holism-reductionism debate. In the competition
37
4 Multifaceted Ecology Between Organicism, Emergentism and Reductionism
over the epistemological determination of ecology, individualistic (Gleason),
anti-organicist and anti-emergentist (Tansley) supporters were ranged against the
upholders of organicist holism (Clements, Phillips; Bergandi 1999; see also Chap. 5).
In the search for the fundamental units of nature, plant ecology played a role of
utmost importance. Various units succeed each other: the biome, the climax, plant
associations and the biotic community. According to Clements (1916) the climax
formation is an organic entity. The formation grows, develops and dies as an organ-
ism. Later, Phillips, following his committed organicist position (1931, 1934,
1935a, 1935b), was to consider the same biotic community as an organism. The
analogy between the organism and the unit of vegetation, the formation or the biotic
community enabled Clements and Phillips to extrapolate certain characteristics
from the first element to the second, albeit with the risk of transforming a relative
similarity into an identity relationship for all aspects – in doing so, there is always
the danger of running into an intellectual dead end. While we have never seen an
organism grow younger, an environmental modification (soil desegregation, for
example) can determine an ecological regression, in other words, species impover-
ishment. However, the phenomenological reading of ecological organicism hides a
more fundamental level of interpretation. These authors, in reality, wanted to point
out the holistic ontological dimension of ecological entities. In other words, they
sought to underscore the “organisational” idea that is inherent in biotic entities.
From this point of view, the influence of philosophical organicism is not to be
completely ruled out. It is interesting to note that the organicist and emergentist
philosophical works of Herbert Spencer, Alfred N. Whitehead, Samuel Alexander,
Conwy L. Morgan and Jan Smuts are all quoted by Phillips and Clements, even if in
relatively late papers (Phillips 1931, 1935b; Clements 1935: see Bergandi 1999).
Moreover, it is noteworthy that Forbes on the one hand and Clements and
Phillips on the other support different forms of organicism. Forbes supports a con-
ception of a biotic community that, while certainly holistic and expressed in organi-
cist terms, is substantially pre-emergentist. His analysis stresses the interactional
dimension between organisms and between organism and environment, whereas
Clements and Phillips are proponents of an organicist perspective which clearly
involves the idea of emergence. For instance, Clements emphasises not only that:
(1) a plant formation is of itself an organism; (2) the climax is the mature stage of the
formation; but also that (3) “the reaction of a community is usually more than
the sum of the reactions of the component species and individuals”, in the sense
that the community naturally produces a cumulative amelioration of the habitat that
would not be possible without the combined action of the individual plants belonging
to the group (1916, pp. 3, 79, 106).
By contrast, Tansley, in a highly paradoxical way, departed from Clements and
Phillips’ organicist perspective by proposing the “ecosystem” concept, which
revealed itself to be a more integrative, holistic entity – the physical system consti-
tuted by the organisms and physical factors. But this proposition neither involved
the disappearance of the other proposed units nor, in its refusal of organicism, was
it able definitively to overcome this epistemological framework (on the definitions
of ecological units, see Jax et al. 1998; Jax 2006). In fact, Tansley identified the
38 D. Bergandi
ecosystem as a “quasi-organism” (1935). In reality, what was at stake was not only
the potentially misleading use of the word “organism”, but above all the principle’s
unpredictability as implied in the organicist community worldview of Clements and
Phillips (Tansley 1935, pp. 297–298). According to Tansley, even if the community
is composed of organisms in mutual association, examination of this entity must be
conducted using an analytic and anti-emergentist perspective. The Tansley refusal
of the Clementsian worldview followed in the footsteps of Gleason’s refusal.
Gleason (1917, 1926) maintained an atomistic and individualistic point of view on
plant association. The lack of limits and structure of the associations was the fun-
damental reason that pushed him to see in these ecological entities the result of
random immigration and environmental variations. This unoriented, random juxta-
position of plants determined structurally different forms of associations, and that
required the total acceptance of analysis as a direct methodological consequence.
Organisms and populations were studied separately, and their association was
reducible to the various isolated plant functions.
The Tansleyan ecosystem concept has had a decisive influence upon successive
phases of the development of ecology. His categorisation of the “basic unit of
nature” was later to be rendered dynamic thanks to Lindeman’s energetic thermo-
dynamics approach (1942), an analytical and additive method that explained the
ecosystem in terms of energy exchanges among the different compartments in the
biotic community and between the community and the physical environment.
Between the 1950s and 1960s, the Odum brothers developed an ecological para-
digm that combines this energetic ecosystem framework with a holistic and emer-
gentist ontology (1953, 1959, 1971: see also 1983, 1993).
The following phrase clearly sums up Eugene Pleasants Odum’s ontological,
methodological and epistemological assumptions.
Just as the properties of water are not predictable if we know only the properties of hydro-
gen and oxygen, so the characteristics of ecosystems cannot be predicted from knowledge
of isolated populations; one must study the forest (i.e., the whole) as well as the trees (i.e.,
the parts). Feibleman (1954) has called this important generalization the ‘theory of integra-
tive levels’. (1971, pp. 5–6)
In other words, ecosystems are complex entities characterised by emergent properties
that cannot be predicted by strictly applying the analytical method. At the same
time, Odum considers his ecology to be the true expression of a holistic approach:
“Practice has caught up with theory in ecology. The holistic approach and ecosys-
tem theory, as emphasized in the first two editions of this book, are now matters of
world-wide concern.” (Odum 1971, p. VII). The issue here is the following: the
Odumian holistic approach takes into account “the ecosystem as a whole”; but
what, precisely, is this “whole”? Is it a matter of ecology, physics or some other
scientific discipline?
In addition, it is interesting to note that Odum considers that “the findings at any
one level aid in the study of another level, but never completely explain the phe-
nomena occurring at that level” (1959, p. 7; 1971, p. 5). Having said this, Odum
seems to deny any value of epistemological reductionism, considering that ecosys-
tem ecology is not reducible to physics. At the same time, it is a matter of fact that
394 Multifaceted Ecology Between Organicism, Emergentism and Reductionism
Eugene Paul Odum, collaborating with his brother Howard Thomas Odum, locates
the theoretical core of systems ecology in energetic analysis:
In ecology, we are fundamentally concerned with the manner in which light is related to
ecological systems, and with the manner in which energy is transformed within the system.
Thus, the relationships between producer plants and consumer animals, between predator
and prey, not to mention the numbers and kinds of organisms in a given environment, are
all limited and controlled by the same basic laws which govern nonliving systems, such as
electric motors or automobiles. (1971, p. 37; see also Chap. 18)
The Odums’ epistemological manifesto has been so effective that from then
onwards ecology has been perceived and presented as the holistic science par excel-
lence.7 In referring to a philosopher of science, Jerome K. Feibleman, they outline
a hierarchical worldview where every level of organisation is characterised by a
specific degree of complexity and properties that are not predictable or explicable
from the study of the lower levels alone (epistemological holism). Implicitly in their
early works and explicitly in the later ones (Odum 1993), the emergence concept and
an emergentist ontology are the cornerstones of the Odumian ecosystem paradigm.
Methodologically, however, they ran into an incoherence that unbalances their
whole theoretical edifice. There are three reasons for this. First, the difference
between collective and emergent properties escapes the Odum brothers, at least in
their early work. Some population and community properties (density, age distribu-
tion, natality, mortality, species diversity, etc.) – even if expressed as statistical
functions – are considered as unique characteristics of the group. In all these cases
the properties, even if they must be considered as group statistical functions, are
determined through the examination of the components using classical analytical
and additive methodology (Salt 1979). Second, the physicalist background of the
Odums’ systems ecology stands in contradiction to emergentist ontological
assumptions. For instance, they consider the outcome of the Eniwetok Atoll energy
evaluation (Odum and Odum 1955; Odum 1977) to be an emergent property. Thus,
they are considering ecological systems as structured physical entities, forgetting
that their specificities are not reducible to the physical domain. Finally, a true emer-
gentist approach will be necessarily a multi-level triadic approach that considers
simultaneously at least the lower and higher adjacent levels in addition to the level
at which the main object of research is to be studied phenomenologically. Instead,
in the Odums’ work the affirmed importance of the emergent properties of ecologi-
cal systems is not coupled with a corresponding emergentist methodology — at
least not until Odum and Barrett (2005, p. 8), where the necessity of a genuinely
triadic emergentist methodology can be clearly recognised. However, the previous
Odumian approach is fully legitimate (see the article by Chap. 15). It is nevertheless
a kind of crypto-reductionist systemism or, to put it in oxymoronic terms, a kind of
reductionist holism, that can at best be considered as “holological” (Hutchinson
1943), and not as the true expression of a holistic, emergentist methodology and
epistemology. Hutchinson proposes making the distinction between holological and
7 The term “holism” was to appear from the third edition (1971) onwards, even if the corresponding
worldview had already been outlined in previous works.
40 D. Bergandi
merological approaches. In a system investigated with a holological approach “(...)
matter and energy changes across its boundaries are studied”, whereas with a mero-
logical approach “(...) the behavior of individual systems of lower order composing
(the system) S are studied” (1943, p. 152). However, it is worth noting that the holo-
logical approach is an expression of a systemic and yet physicalist perspective,
while the mereological one, methodologically speaking, is a strict expression of an
analytical-additive reductionist perspective. McIntosh (1985, pp. 199–213), Taylor
and Blum (1991, p. 284; see also Taylor 2005) were among the first to analyse the
Janus-like character of the ecosystem ecology represented by E.P. Odum: they saw it
as a “functionally holistic” new ecology, which was essentially, however, expressed
through system modelling involving the physical attributes of ecosystems.
Conclusion
The holism-reductionism debate in ecology is, without a doubt, a protean issue. In
ecological studies, first, the debate took a number of forms: an organicist world-
view that expressed the holistic, systemic relations existing between organisms, and
between organisms and their environment (Forbes); an organicist and emergentist
view of plant communities (Clements, Phillips); a view of plant associations as
individualistic, atomistic, randomly generated entities (Gleason); and, finally,
Tansley’s integrative “ecosystem” concept that expressed the epistemological
refusal of Clementsian organicism and emergentism.
Second, it showed itself in the form of the acceptance or refusal of physicalism.
If we broach the epistemological nature of ecology and come to the conclusion that
ecology is fundamentally a holistic science, we would be mistaken in thinking that,
methodologically, ecology necessarily embodies an emergentist approach. In fact,
an “emergentist holistic” approach need be understood neither as a reiteration nor
as a tautology. This distinction is of utmost importance. It enables us to avoid all
the inconsistencies inherent in the Odumian paradigm and all paradigms that pro-
pose a holistic ontology but that, in practical methodological research, deploy the
full panoply of reductionism. In fact, in the history of science, a holistic and emer-
gentist ontology is not always applied consistently in emergentist methodology and
epistemology. For instance, once we cease to consider ecosystem ecology as the
expression of a “holistic” attitude and recognise in it instead a “holological” frame-
work, a kind of oxymoronic “reductionist holism”, then we will avoid misunder-
standings and be back on track. We will then be free to construct a truly consistent
holistic and emergentist ontological, methodological and epistemological
framework.
Finally, to sum up from a strictly epistemological point of view, one of the major
implications of the holism-reductionism debate is the confrontation between two
philosophies that at first sight support a shared hierarchical worldview of natural
reality. However, there is one very important difference. From the reductionist point
of view, the ideal point to reach is that all the scientific disciplines will, sooner or
later, be formulated, interpreted and reduced to the more fundamental sciences,
414 Multifaceted Ecology Between Organicism, Emergentism and Reductionism
particularly physics. From the holistic or, more correctly, emergentist point of view,
the supposed ontological natural hierarchy does not involve a hierarchical relation-
ship between the scientific disciplines but rather a systemic one. The sciences with
their specificities and particularities allow us to grasp different aspects of reality
which we cannot reduce to one another, but which we can combine in order to
arrive at a non-definitive, ever-changing picture of reality.
For emergentists, the universe is a growing entity that generates ever new phe-
nomena, events and qualities which can be neither predicted nor deduced from
those that preceded them. We must remember, however, that an a priori unpredict-
ability does not necessarily involve the refusal of an (ideal) a posteriori explanation.
On the contrary, according to anti-emergentists “nothing is new under the sun” and,
above all, any so-called novelty is predictable and explicable: the same universe,
yet two antinomic and incommensurable worldviews. Which paradigm is closer to
reality? Are the reductionists correct when they claim emergents are epiphenom-
ena? Are the emergentists wrong when they attribute an ontological status to emer-
gence and, above all, axiomatically assert its a priori unpredictability? These are
open questions to which the answers will probably never be given once and for all
but always case by case.
Finally, to grasp the logical structure of alleged emergence, we must ask our-
selves: what are the emergents – properties, relations, entities or laws? What is the
level of organisation that bears the property which is supposed to be emergent? In
addition, the levels of organisation – and among them, significantly, certain eco-
logical levels such as the ecosystem or landscape – must be understood as levels of
description, or “methodological abstractions”. These epistemological fictions
sometimes make it possible to develop models that allow us to approach natural
reality “asymptotically”. Otherwise we risk an insidious epistemological fallacy: a
hypostatisation of our abstractions that brings us to project our hypotheses and
theories onto reality, forgetting that they are merely notional tools by which to
approach it. A metaphor may help to clarify this idea: it is like a dog that starts to
play with you but forgets, in the excitement of the game, that it is playing and
begins to bite in earnest. In other words, a constructivist epistemology prevents us
from being bitten by the rock-hard certitudes of naive realism. Our scientific con-
structs make it possible to approach natural reality without ever fully grasping it.
These constructs allow us to understand certain aspects of reality in a non-definitive
way. They remain valuable until such time as new constructs allow us to get even
closer. This is a genuine process of scientific knowledge where the syntagm “The
End” will never be written.
References
Ayala FJ (1974) Introduction. In: Ayala FJ, Dobzhansky T (eds) Studies in the philosophy of
biology. Reduction and related problems. MacMillan, London, pp vii–xvi
Beckermann A, Flor H, Kim J (1992) Emergence or reduction? De Gruyter, Berlin
Bergandi D (1995) ‘Reductionist holism’: an oxymoron or a philosophical chimaera of
E.P. Odum’s systems ecology. Ludus Vitalis, 3, 5, pp 145–180; reprinted in Keller DR, Golley
42 D. Bergandi
FB (eds) (2000) The philosophy of ecology: from science to synthesis. University of
Georgia, Athens (abridged version), pp 204–217
Bergandi D (1999) Les métamorphoses de l’organicisme en écologie: de la communauté végétale
aux ecosystems. Revue d’histoire des sciences 52(1):5–31
Bergandi D (2007) Niveaux d’organisation: évolution, écologie et transaction. In: Martin T (ed)
Le tout et les parties dans les systèmes naturels. Vuibert, Paris
Bunge M (1991) The power and limits of reduction. In: Agazzi E (ed) The problem of reduction-
ism in science. Kluwer, Dordrecht, pp 31–49
Campbell DT (1974) ‘Downward causation’ in hierarchically organized biological systems. In:
Ayala FJ, Dobzhansky T (eds) Studies in the philosophy of biology. MacMillan, London, pp
179–186
Clements FE (1916) Plant succession: an analysis of the development of vegetation. Carnegie
Institution, Washington, DC, p 242
Clements FE (1935) Experimental ecology in the public service. Ecology 16:342–363
Dewey J, Bentley AF (1949) Knowing and the known. Beacon, Boston
Duhem P (1977) The aim and structure of physical theory. Atheneum, New York
Elton CS (1927) Animal ecology. Sidgwick & Jackson, London
El-Hani CN, Pereira AM (2000) Higher-level descriptions: why should we preserve them? In:
Andersen PB, Emmeche C, Finnemann NO, Christiansen PV (eds) Downward causation:
minds, bodies and matter. Aarhus University Press, Aarhus, pp 118–142
Feibleman JK (1954) Theory of integrative levels. Br J Philos Sci 5:59–66
Forbes SA (1880) On some interactions of organisms. Ill Nat Hist Surv Bull 1(3):3–17
Forbes SA (1887) The lake as a microcosm. Ill Nat Hist Surv Bull 15(9):537–550
Forbes SA (1914) Fresh water fishes and their ecology. Illinois State Laboratory of Natural
History, Urbana (read at the University of Chicago, August 20, 1913)
Gleason HA (1917) The structure and development of the plant association. Bull Torrey Bot Club
44:411–462
Gleason HA (1926) The individualistic concept of the plant association. Bull Torrey Bot Club
53:7–26
Haeckel E (1866) Generelle Morphologie der Organismen. Allgemeine Grundzüge der organis-
chen Formen-Wissenschaft, mechanisch begründet durch die von Charles Darwin reformirte
Descendenz-Theorie. Reimer, Berlin
Hempel CG, Oppenheim P (1948) Studies in the logic of explanation. Philos Sci 15:135–157
Hutchinson GE (1943) Food, time, and culture. N Y Acad Sci 15:152–154
Jax K (2006) Ecological units: definitions and application. Q Rev Biol 81(3):237–258
Jax K, Jones CG, Pickett STA (1998) The self-identity of ecological units. Oikos 82(2):253–264
Jones R (2000) Reductionism: analysis and the fullness of reality. Bucknell University,
Lewisburg
Levins R, Lewontin R (1980) Dialetics and reductionism in ecology. In: Saarinen E (ed)
Conceptual issues in ecology. D. Reidel, Dordrecht, pp 107–138
Lindeman RL (1942) The trophic-dynamic aspect of ecology. Ecology 23:399–418
McIntosh RP (1985) The background of ecology. Concept and theory. Cambridge University
Press, Cambridge
Mayr E (1988) Toward a new philosophy of biology. Belknap/Harvard University Press,
Cambridge
Nagel E (1961) The structure of science: problems in the logic of scientific explanation. Brace and
World, New York, Harcourt
Odum EP (1953) Fundamentals of ecology. W.B. Saunders, Philadelphia
Odum EP (1959) Fundamentals of ecology. W.B. Saunders, Philadelphia
Odum EP (1971) Fundamentals of ecology. W.B. Saunders, Philadelphia
Odum EP (1977) The emergence of ecology as a new integrative discipline. Science 195:
1289–1293
Odum EP (1983) Basic ecology. W.B. Saunders, Philadelphia
Odum EP (1993) Ecology and our endangered life-support systems. Sinauer, Sunderland
434 Multifaceted Ecology Between Organicism, Emergentism and Reductionism
Odum EP, Barrett GW (2005) Fundamentals of ecology, 5th edn. Thomson brooks, Belmont
Odum HT, Odum EP (1955) Trophic structure and productivity of a windward coral reef com-
munity on Eniwetok Atoll. Ecol Monogr 25:291–320
Phillips J (1931) The biotic community. J Ecol 19:1–24
Phillips J (1934) Succession, development, the climax and the complex organism: an analysis of
concept. J Ecol 22(1):554–571
Phillips J (1935a) Succession, development, the climax and the complex organism: an analysis of
concept. J Ecol 23(2):210–246
Phillips J (1935b) Succession, development, the climax and the complex organism: an analysis of
concept. J Ecol 23(3):488–508
Popper KR (1972) Objective knowledge: an evolutionary approach. Clarendon, Oxford
Quine VOW (1961) From a logical point of view. Harvard University Press, Cambridge
Ruse M (1988) Philosophy of biology today. State University of New York, Albany
Salt GW (1979) A comment on the use of the term emergent properties. Am Nat 113:145–149
Salthe SN (1985) Evolving hierarchical systems: Their structure and representation. Columbia
University Press, New York
Schwarz AE (2003) Wasserwüste – Mikrokosmos – Ökosystem. Eine Geschichte der Eroberung
des Wasserraumes. Rombach-Verlag, Freiburg
Tansley AG (1935) The use and abuse of vegetational concepts and termes. Ecology
16(3):284–307
Taylor PJ (2005) Unruly complexity: ecology, interpretation, engagement. The University of
Chicago, Chicago
Taylor PJ, Blum AS (1991) Ecosystem as circuits: diagrams and the limits of physical analogies.
Biol Philos 6:275–294
Whitehead AN (1925) Science in the modern world. MacMillan, New York
Williams GC (1966) Adaptation and natural selection: a critique of some current evolutionary
thought. Princeton University Press, Princeton, New Jersey
Williams GC (1985) A defense of reductionism in evolutionary biology. In: Dawkins R, Ridley M
(eds) Oxford surveys in evolutionary biology, vol 2. Oxford University Press, Oxford, pp 1–27
Woodger JH (1952) Biology and language. Cambridge University Press, Cambridge
45
Introduction
Controversies between holism and reductionism are a familiar feature in many fields
of inquiry besides ecology. Although seldom described in these terms,1 the issue has
long played – and continues to play – a significant role in science, philosophy and
political ideology; indeed, one could almost say that it has always existed. It played
a major part, for example, in shaping the ideological conflicts between conservatism
and liberalism in the nineteenth and twentieth centuries. Holistic ideas were present
especially in “Lebensphilosophie” (philosophy of life) and in other philosophies
critical of science, such as historicism, that shaped the zeitgeist at the time of eco-
logy’s emergence. Towards the end of the twentieth century, holism became a sig-
nificant force in “political ecology”. This is true not only of those strands of political
ecology and related fields that are explicitly committed to the task of “renewing” the
world view of their time, such as “Deep Ecology” (Naess 1973; Drengson and Inoue
1995) and the “New Age Movement” (Capra 1982); rather, the view of nature within
all political ecology is essentially a holistic one. However, a range of conceptual
figures associated with holism can be found in many older philosophies as well. The
macro-microcosm figure, for example, appears as far back as Plato (cf. Schwarz
2003) and came to exert influence within modernity mainly through the world view
that underlies Leibnizian rationalism (cf. Eisel 1991; Langthaler 1992). Nowadays
research programmes in most of the natural sciences are shaped by reductionist ideas;
they are linked primarily to neo-positivist philosophies and can be traced back to
older empiricist philosophies, as well as to Cartesian rationalism.
The holism-reductionism debate in ecology can be properly understood only
against this non-scientific backdrop, because what is at issue in ecology is more than
Chapter 5
The Classical Holism-Reductionism
Debate in Ecology
Ludwig Trepl and Annette Voigt
1 The term “reductionism”, for example, became common only in the middle of the twentieth cen-
tury, even though it covers older philosophical problems, which previously came under the rubric
of “materialism” or “mechanism”, and the methodology of specific sciences (cf. Stöckler 1992).
A. Voigt (*)
Urban and Landscape Ecology Group, University of Salzburg, Hellbrunnerstraße 34,
5020 Salzburg, Austria
e-mail: annette.voigt@sbg.ac.at
A. Schwarz and K. Jax (eds.), Ecology Revisited: Reflecting on Concepts, Advancing Science,
DOI 10.1007/978-90-481-9744-6_5, © Springer Science+Business Media B.V. 2011
46 L. Trepl and A. Voigt
just whether scientific theories of a certain type provide a correct description of certain
natural phenomena. What these philosophical and political-ideological controversies
demonstrate above all is that the issue is of relevance in areas that lie well beyond the
confines of the discipline itself. It therefore appears justifiable to us to analyse a wide
range of debates on the basis of the holism-reductionism complex, including those that
are not explicitly about it at all.2 The significance of the complex is by no means
exhausted in the impact it has on ecology or on understandings of ecology.
The holism-reductionism controversy in ecology is all about the relationship
between wholes and their parts. This is a major problem in many sciences – in
physiology (the organism), geography (the landscape), psychology (the soul) and
sociology (society), as well as in physics, linguistics and epistemology. However,
the debate in ecology is of particular interest when it comes to understanding the
way holistic and reductionist ideas about nature and society work in the context of
political-ideological controversies. The ecological debate has one thing in particu-
lar in common with a very few sciences (with sociology above all, though not with
other natural sciences), something that links it very closely to those ideological
struggles, and that is this: ecology is one of those sciences whose objects of inquiry
are constituted in such a way that the parts are usually individuals and the whole a
community. The contrast between reductionism and holism in ecology tends to take
the form of an opposition between individualism and organicism. In turn, individu-
alism, as a form of reductionism, is holistic in the sense that while individuals are
thought of as wholes (as organisms), a community is not. “Community”, according
to individualism, is merely a name for a certain number of individuals, gathered
together more or less at random by the scientist, who are considered to be “autono-
mous” and who alone are seen as being real. In organicism, by contrast, “commu-
nity” is conceived of as an organic community or as a superorganism, in other
words, the relationship between the part and the whole is conceptualised in analogy
to the relationship between organ and organism. – Whenever organicism or the
“organismic concept” is mentioned in ecology, the reference is to something differ-
ent from what, in biological terms, is commonly called the “organismic approach”
or “organismic biology”. The idea entailed by the former is that a “Lebensgemeinschaft”
(biotic community) is of the same character as an individual organism; in the latter,
importance is generally attached instead to the level of the individual organism, the
point here being to counter the tendency to focus solely on the molecular level.
In the following we shall refer to only a few of the many variants of the holism-
reductionism debate in ecology and to certain stages of their transformation. Our
main concern is to reconstruct the logic of the debate and to determine the concep-
tual structures on which it is based. Given that the positions adopted address a
problem that not only exists within science but is above all of a fundamental philo-
sophical nature, we can interpret the actual emergence of such positions and their
transformation in ecology in the following way. There are certain ways of concep-
tualising the relationship between parts and wholes, or between individuals and
community – there is not an indefinite number of variants of these conceptual
2 Cf. Trepl (1994) and the critical response by Levins and Lewontin (1994).
475 The Classical Holism-Reductionism Debate in Ecology
figures. What we are interested in are the conditions in which certain combinations
of their elements and certain transformations in their structure are possible; above
all, we are interested in the practical and ideological implications of these concep-
tual figures. We do not intend to present every single “important” theory that has
ever made an appearance in the history of ecology; instead, our aim is to construct
ideal types. These enable us to present the positions that were actually adopted
throughout history and to compare them in a systematic way. Our main criterion for
selecting the examples is less that they are considered, for whatever reasons, to be
important in ecology – the fact that they have been influential, for example – but
rather that they are suitable for explicating the ideal type constructions.
Numerous potential variants of holism and reductionism exist and can be found in
biology. We mention this briefly at the start. The main point here is that the spectrum
of potential variants is by no means exhausted by organicism and individualism. Than,
we present reconstructions of ideal types of each in its classical form as they emerged
during the first few decades of the twentieth century, using examples by way of illus-
tration. Since the dispute between them cannot be resolved empirically, we inquire as
to whether this might be possible at the methodological level. It proves to be difficult
at this level as well. We choose an approach based on a theory of constitution,3 which
enables us to regard both holism and reductionism as being “inspired” by certain
world views. On this basis it becomes easier to understand the utterly different practi-
cal consequences entailed by holistic ecological theories on the one hand and reduc-
tionist theories on the other. We conclude with an example that describes the dynamic
through which both approaches (usually in response to one another) change.
Variants of Holism and Reductionism
The literature refers to holism and reductionism in many very different ways.4 In the
following we take a few examples and explain briefly how they work conceptually
on the basis of what is common to all forms of holism and reductionism. Our
examples are restricted to those variants that play a role in biology, so they are about
explaining “life”.
3 Constitution here is understood differently from the sense common in philosophy, that is, the way
Kant, for example, used it. Instead, it refers to the idea that scientific theories are not simply genera-
lised depictions of specific empirical observations but that they owe their existence to non-scientific
conceptual structures already in existence. Thus, we can speak here of a “l´a priorí historique” (his-
torical apriori) (Foucault 1969), of “bereitliegenden kulturellen Deutungsmustern” (cultural patterns
of interpretation that are already available) or of “Konstitutionsideen” (ideas of constitution) (Eisel
2002, p. 130), which are the conditions of possibility – realized through culture – for scientific con-
cepts and for their corresponding objective experiences. Indeed it is these conditions which ensure
that new facts do not destroy the old conceptions in general but consistently confirm the theory (or
the paradigm) (Eisel 2002; cf. Kuhn 1962).
4 Some examples of literature that takes this issue further include: (science in general) Nagel 1949,
1961; Bueno 1990; Agazzi 1991; (biology) Ayala 1974; Ayala and Dobzhansky 1974; Ruse 1973;
Hull and Ruse 1998; Bock and Goode 1998; Looijen 2000; (ecology specifically) Saarinen 1982;
Bergandi 1995; Bergandi and Blandin 1998; Keller and Golley 2000; Kirchhoff 2007; Voigt 2009.
48 L. Trepl and A. Voigt
It is probably true to say that what links all those things together that are referred
to as holism is not much more than the principle that the “whole” has “priority”
over the “parts” – whatever “priority” might mean exactly – and a set of reserva-
tions about any form of “simplification”. On the reductionist side, the commonality
between different positions probably consists above all in their emphasising that
statements about phenomena of a complex nature should be derived from state-
ments about phenomena of a simpler nature, and that science essentially consists in
this kind of “reduction”.
Different forms of holism and reductionism also come about depending on the
aspects of the research object (or of the scientist’s relationship to that object) to
which these principles are applied. In other words, it is not only certain methods or
research programmes that can be called holistic or reductionist, but also certain
views about the “nature” of their objects. This means that whether a certain position
appears to be holistic or not depends on the perspective taken. We shall discuss here
just a few of the numerous permutations possible: “wholeness” can be taken to
mean a variety of very different things (2.1); “simplification” can mean very differ-
ent things (2.2); the assertion that something is reductionistic or holistic may be a
reference – among other things – to the nature of reality or to how we should pro-
ceed in order to find something out about it (2.3).5
Aspects of the Concept of “Wholeness”
Those methods and theories that are termed holistic differ greatly according to
which aspect of the concept of wholeness they highlight. In biology, such aspects
include totality, gestalt, uniqueness, system character and “Lebendigkeit” (alive-
ness) – although often many of these aspects cannot be separated from one
another.
In the case of “aliveness”, the choice to focus on one or the other aspect has far-
reaching methodological consequences. For example, the whole can be identified
as an “inner essence” (e.g. “soul”); one point of access to this wholeness can be
seen as being that the relationship between the inside, which remains hidden, and
the outside, which is perceived, is one of the latter giving expression to the former.
It then becomes possible to draw on methodologies from the human sciences whose
aim is to “understand” this inner essence through its representation in the external
world (especially Dilthey’s “Ausdrucksverstehen” around the turn of the twentieth
5 Other levels in addition to the ontological and methodological would be the epistemological level
(“Erkenntnistheorie”), which is about the validity of knowledge, and the level of theory of science
in the narrow sense (“Wissenschaftstheorie im engeren Sinne”), which is about the character of the
empirical phenomenon of science. One might also add a level of theory of constitution, at which
the independence of the ontological and the epistemological level disappears. One could, for
example, describe the theory of Thomas Kuhn as on the level of theory of science holism in the
narrower sense mentioned. Coherence theories of truth (e.g. the Duhem-Quine theory) might be
called epistemological holism (cf. e.g. Oppenheim and Putnam 1958; Ayala 1974; Putnam 1987).
495 The Classical Holism-Reductionism Debate in Ecology
century, cf. Dilthey 1883). External forms are largely understood in terms of a
gestalt. Even if every view of the whole as a gestalt does not imply such a relation-
ship of representation, this is nonetheless very often the case. Examples include
work by Portmann (e.g. 1948) and Troll (e.g. Wolf and Troll 1940; Troll 1941), who
refer to the morphology of individual organisms. But even in relation to objects
such as vegetation there are “physiognomic” approaches that correspond to this
model. Of particular significance, historically speaking, is Grisebach’s (1838) con-
cept of formation, whose relationship to Humboldt’s “Physiognomics of Plant Life”
(Humboldt 1806) clearly places its origins in an approach based firmly on
“Ausdrucksverstehen” (cf. Trepl 1987, pp. 103 ff.). The gestalt aspects of this are
usually linked to other aspects of wholeness, such as the organic interaction of
parts. Holistic positions of this kind refuse to conform – sometimes avowedly so – to
scientific demands, insofar as they counter the latter with a “a vivid and clear
idea”,6 a holistic “Naturschau” (contemplation of nature, Thienemann 1954, p. 322)
or a “contemplative look at nature of a morphological kind”7 and declare these to
be the goal of biology.8
Despite having what is, in principle, a similar conception of inner essence as a
wholeness, others claim not to depart at all from the scientific methodological ideal.
In neovitalism, for example, the specificity of life was seen in it being characterized
by a “life force” (entelechy). This is not regarded as a physical force and certainly
not one that can be measured scientifically; instead, it is seen as being a living,
soul-like force, which is gestalt-forming and therefore holistic. Despite this, ent-
elechy (known as “Factor E”) is claimed to be “empirically real”.9 Vitalism has
been accused of being dualistic by authors who have been described as holists in
the history of biology (e.g. Bertalanffy, Haldane). Vitalism, they say, sees in the
living organism only a sum of so many parts that are complemented and monitored
by a kind of soul in the role of engineer, rather than seeing the essence of life in the
interactive structure of the whole (Bertalanffy 1949, p. 30). It is this, namely, the
organic interaction of the parts, that constitutes the holistic element in life. In this
view, biological holism – that is, what was explicitly known as holism in the history
of biology and in philosophies related to biology (the views of J. S. Haldane, Smuts
and A. Meyer-Abich, for example), as well as in systems theories with a holistic
orientation in the tradition of Bertalanffy (see also Chap. 15) – is given when the
key element of life is not seen to lie in an inner force inaccessible to scientific
methods. Biological holism consists, instead, in the view that the characteristic of
being alive can only be attributed to objects that are a whole, and that this whole
exists in a special relationship to its parts that is not found in non-living objects.
These wholes, so the theory goes, require an approach of their own that is different
from that of physics. To the extent that holistic theories divide reality into different
6 “bildhafte, anschauliche Vorstellung” (Friederichs 1957, p. 120, cf. also 1937).
7 “anschauende Naturbetrachtung morphologischer Art” (Thienemann 1954, p. 317).
8 Cf. on biology as a whole: Köchy 1997, 2003; cf. on ecology: Trepl 1987; Jax 1998, 2002.
9 “empirisch wirklich” (Driesch 1935, p. 75, cf. also Mocek 1974, 1998).
50 L. Trepl and A. Voigt
levels or autonomous wholes to which different scientific methods have to be
applied respectively, they can be called pluralistic.
Different Kinds of Reduction
Reductionism is used to refer to a situation in which, during the course of develop-
ment of a science, a requirement is made of all its theories that they should be based
on the theories of a basic science. In biology, reduction is understood principally in
terms of tracing back something that is living to something that is not living by
means of a physical-chemical explanation of specific metabolic processes, for
example. Reductionism essentially coincides with what is often called “mechani-
cism” (or “mechanism”) or else “physicalism”.10
Two forms need to be distinguished here in particular:
1. Some hold the view that the whole needs to be explained by acquiring knowledge
about its parts (a “bottom up” approach). However, reduction is deemed to have
been successful only when these parts have been reduced to certain “things”,
namely “fundamental units” (“atomistic” reductionism). These fundamental units
are not alive; in other words, this form of reductionism assumes that even if the
whole is an organism, there is always a subordinate level in the hierarchy (in the
sense of a “nested hierarchy”) at which the parts can no longer be considered to
be living (subcellular level, molecular level). However, by conducting research on
these, it is possible to obtain full knowledge of the whole, the living organism.
Examples are superfluous here, as this approach constitutes the mainstream in
biology. Indeed it was this form of reductionism, known as “mechanism” (cf. e.g.
Roux 1895; Loeb 1916), which was discussed most among biologists during the
period when ecology was emerging: the phenomena that could be observed in
organisms could ultimately be explained causally at the molecular level. Both
Darwinism and neo-Darwinism were also described in terms of mechanistic
reductionism. In this case, however, the individual organism is not reduced to the
molecular level; instead, in its mechanistic explanations of evolution, Darwinism
always presupposes the organism as a whole. Events occurring at the molecular
level only become relevant in terms of evolutionary biology when they are viewed
in relation to the organism (e.g. as a contribution to its fitness). Evolution is
explained mechanically on the basis of interactions between organisms11 and of
those between organisms and their abiotic environment.
10 The accusation of reductionism means that the simplification is carried out in such a way that
it leads precisely to not explaining, or explaining wrongly, the matter to be explained, such as a
living organism.
11 The fact that in the context of Darwinism other levels – the individual gene or the population – are
shifted to centre stage changes nothing of this fundamentally organism-centred structure of
Darwinism. Even if individual genes are taken as a starting point, they nonetheless “want” some-
thing, as “selfish” genes (Dawkins 1976) and are not simply chemical-physical phenomena.
515 The Classical Holism-Reductionism Debate in Ecology
2. It is also possible to undertake a physical-chemical reduction quite independently
of the issue of levels in a “nested hierarchy”. One example of such a reduction is
the “physicalisation” of the organism, for example by looking at blood circulation
as a hydraulic system (Harvey in the seventeenth century). Unlike reduction to the
molecular level, the whole here is reduced primarily not to its parts but to pro-
cesses or characteristics of all its parts (e.g. flow speed). Measuring these is seen
as a way of comprehending the whole. Parts of the system are addressed from a
common functional perspective, where the function often lies in contributing
towards maintaining particular processes. It is thus possible for an extreme form
of reductionism to appear, from a different perspective, to be holism12 insofar, for
example, as everything about an object is expressed in energetic terms.
Methodological and Ontological Holism/Reductionism
According to methodological reductionism, traditional (or typically) biological
explanations should be replaced by physical-chemical ones, i.e. functional explana-
tions should, in general, be replaced by causal ones. This generates the possibility
of explaining biological phenomena in a strictly scientific way.13 However, this says
nothing about the way of being (“Seinsweise”) of the object in question, because
all that is being argued is that it seems advisable to undertake such a reduction on
methodological grounds (e.g. the call for the greatest possible simplicity of the
entire body of theory). Methodological reductionism here is diametrically opposed
to methodological holism,14 which insists that the specific biological mode of
explanation should be retained. Assuming we are dealing with methodological
holism only, the claim being made is not that specifically biological terms (e.g.
maturity, stimulus, mating instinct) describe objective facts which cannot be
captured in physical-chemical terms, but rather that their use is merely understood
to be methodologically useful or necessary. Even explicitly teleological explana-
tions, which ultimately always refer to the phenomenon to be explained in terms of
its significance in the context of a whole, are permissible in this sense, because
without them we would have difficulty – indeed, it would be impossible – to ask
questions that are relevant in view of the specific phenomena with which biology
is concerned.15 Thus, teleological explanations are of heuristic value – in fact,
12 “The thermodynamic approach is particularly well fitted as a tool to describe ecosystems from
an holistic point of view because it is based on the macroscopic flows of energy and mass”
(Jørgensen 2000, p. 113).
13 On whether and to what extent functional explanations are also teleological, see e.g. Nagel
(1979); Rosenberg (1985); Mayr (1988); McLaughlin (2001).
14 “Methodological holism” is often also used to describe a social scientific view according to
which social relations can only be interpreted and explained in terms of social wholes (e.g. classes,
but especially the society as a whole) (Mittelstraß 1995, p. 123).
15 Cf. especially Cassirer (1921), as well as the whole tradition of meta-theory in biology based on
Kant; it can generally be seen as a form of methodological holism.
52 L. Trepl and A. Voigt
in this respect, they are indispensable. This kind of methodological holism is
compatible with ontological reductionism, just as methodological reductionism is com-
patible with ontological holism (cf. also Mayr 1982).
According to ontological reductionism, everything that exists consists of “funda-
mental” elements: “an organism is essentially nothing but a collection of atoms and
molecules” (Crick 1966).16 This assumption about the character of being implies that
the characteristics of higher forms of organisation can generally be explained (com-
pletely) causally as being due to the mutual influence of fundamental elements
(“micro-determination”). This also provides the grounds for limiting the types of
terminology allowed in science. Biology can be expressed in terms of physics and
chemistry. Contrary to this position we have a form of ontological holism whose
core assertion (as far as biology is concerned) is this: organisms are not the way they
appear in their reductionist physical-chemical guise. It is generally assumed that
units found at higher levels of organisation are comprised of nothing other than units
at lower levels (that is, for example, organisms are made up of organs, the latter of
cells, and cells of molecules); there is no suggestion here that there is also a vital
force at work. However, the complexity and organisational structure of such higher-
level units make it impossible to say that they are “nothing but” a collection of
“fundamental” units. A hierarchical order of increasing complexity can, it is said, be
set up (cf. the familiar representations of life as a “layered structure”: atom, mole-
cule, organelle, cell, tissue, organ, organism, community etc.). At every higher level
of organisation, one finds emergent, or irreducible, characteristics – hence the insis-
tence on the autonomy of biology. But as far as methodology is concerned, it is
perfectly possible in this context to hold the view that a physical-chemical reduction
is useful or even necessary, namely as a connecting theme for research (method-
ological reductionism).17 It is just that this will not be linked to the view that higher-
level units can be explained in their entirety and that biology will ultimately prove
to be explicable in terms of physics and chemistry: life is essentially something other
than the objects that are accessible to these sciences. This ontological holism, then,
could be compatible with methodological reductionism (see Putnam 1987). In the
context of ontological holism, though, it is also possible to hold the methodological-
holistic view, mentioned above, that reduction is not an option at all if one wishes
to acquire insight into a living object of nature.
As in biology as a whole, the holism-reductionism debate in ecology is all about
the “special conditions for interpretation and explanation of partial groups of
objects”18 and, in particular, about the question of whether irreducible wholes exist
or not. This controversy acquires a special twist in ecology, however: If reducing a
16 What was described above as “atomistic reductionism” may be such an ontological reductionism,
but it may also be meant merely in the sense of a methodological rule.
17 If we emphasise this and not the heuristic indispensability of teleology, then the Kantian tradi-
tion mentioned above should be described as methodological reductionism.
18 “besonderen Deutungs- und Erklärungsbedingungen partieller Gegenstandsbereiche”
(Mittelstraß 1995, p. 123).
535 The Classical Holism-Reductionism Debate in Ecology
unit consisting of interacting organisms to its parts is not continued to a level at
which the parts are no longer considered to be living, then the relationship between
whole and parts becomes a relationship between community and individuals (both
community and individual being understood in the broadest sense). In its relation-
ship to individuals, community can be conceptualised according to several different
models. Let us look at two of these first of all. Community can be conceptualised:
1. as “organische Gemeinschaft” (organic community), or rather as an organism of
a higher order, so that the parts (individuals) become (components of) organs of
the whole;
2. as a community in the narrower sense (“Gesellschaft” as opposed to
“Gemeinschaft”), i.e. as an interactive network that comes about when individu-
als that are independent in principle enter into relations of cause and effect with
other individuals (or, in borderline cases, when no relations of cause and effect
exist anymore and the community becomes an aggregation of unconnected
individuals).
The form of holism mentioned in (1) above is the one that predominates in
ecology. Whenever ecologists speak of organicism, the view being referred to is the
one that communities of organisms are themselves organised like organisms. In the
context of biology, this is a form of holism specific to ecology, because within
biology communities of organisms are the object of ecological inquiry by defini-
tion. (2) is the form of reductionism typical in ecology, namely individualism. The
model of the machine takes up a mediating position between the two models of
organism and community. The machine is a whole, only it is one that functions not
organically but mechanically, one whose parts (are supposed to) fulfil a function
together. That function does not lie, however, in generating or maintaining itself,
and the whole is not grown but rather constructed. This model is relevant to the
dynamic that arises when the individualistic and organicist approaches encounter
one another and are exposed to different kinds of external conditions. First of all,
however, we are concerned only with the first two models mentioned.
Holism and Reductionism in Classical Ecology
The development of holism in biology – and therefore in ecology as well – is attrib-
uted mainly to the biologists J. S. Haldane, Bertalanffy and Needham,19 as well as
to certain non-biological influences.20 Nonetheless, the role of these authors ought
not to be overestimated, despite the fact that they were embraced by ecologists
themselves and, probably on account of this, were considered in history of science
accounts to be particularly influential. Certain less direct influences were almost
19 E.g. Haldane (1931); Needham (1932); Bertalanffy (1932, 1949).
20 E.g. Smuts (1926); cf. his influences on ecology in Phillips (1934, 1935), Bews (1935); on this,
see e.g. McIntosh (1985); Trepl (1987); Hagen (1992); Anker (2001).
54 L. Trepl and A. Voigt
certainly of much greater significance. When ecology emerged towards the end of
the nineteenth century, and even in the first few decades of the twentieth, the zeit-
geist was strongly influenced by holistic views,21 which were an essential aspect of
conservative philosophies and ideologies of the time that were critical of civilisa-
tion and were anti-mechanistic (cf. Harrington 1996, Müller 1996). Older philoso-
phies also played a not inconsiderable role (e.g. those of Herder and, by way of the
latter, Leibniz; cf. e.g. Eisel 1980, see also Chap. 25), as they were of paradigmatic
significance for the new geography – ecology initially developed in very close
association with physical geography. The paradigmatic core of geography was the
unit of culture and concrete nature, conceived of as an organism and known as
“Land” (land) or “Landschaft” (landscape) (cf. Eisel 1980).
Frequently named as the representatives of an early holistic strand among ecolo-
gists are Friederichs (1927, 1934, 1937), Thienemann (1941, 1944), Thienemann
and Kieffer (1916), Clements (1916, 1936), Shelford (especially Clements and
Shelford (1939)), Phillips (1934, 1935), Braun-Blanquet (1928) and Sukachev
(1958).22 While their views of the essence of “synecological” objects (“biocoenose”,
“community”, “holocoen”) differ in detail, they can still be characterised overall as
organicist in the sense described above.
One important contrasting position to the “renewal” the sciences were undergo-
ing on the basis of different concepts of wholeness was positivism and, later, in the
twentieth century, neo-positivism (logical positivism). To the latter, statements
about wholeness are considered to be metaphysical. This basic attitude is reflected
in ecology – although it owes its existence by no means to the influence of (neo-)
positivism alone, the latter being just one of several philosophies at the time that
were oriented towards the exact sciences. Representatives of this strand include
Forbes (1887), Gleason (1917, 1926, 1927), Ramensky (1926), Lenoble (1926),
Gams (1918) and Peus (1954).23
In the following we reconstruct the classical positions of organicist holism and
reductionist individualism as ideal types. An ideal type does not claim to be an
entirely real position – even though our constructions draw on formulations put
forward in particular by Clements (1916, 1936) on the one hand and Gleason
21 These views did not constitute a single coherent position, but rather very different ones, of which
only a few are (explicitly) conservative and critical of civilisation (certainly not gestalt theory –
Ehrenfels 1890, 1916; Wertheimer 1912; Köhler 1920 – or holistic approaches in neurology, e.g.
Goldstein 1934). The greatest influence was exerted by some authors who were adherents of the
philosophy of life (such as Klages, Spengler, Bergson) and other philosophers (e.g. Husserl,
Heidegger, Whitehead). Although only minimally influential in the public intellectual sphere,
Smuts (1926) and Meyer-Abich (1934, 1948) were highly regarded in biology; the philosophical
and social scientific works of biologists, such as Uexküll’s “Staatsbiologie” (Uexküll 1920) (State
biology) should also be mentioned here.
22 Cf. also McIntosh (1985); Trepl (1987); Botkin (1990); Jax (2002); Kirchhoff 2007; Voigt 2009.
23 On this, see also McIntosh (1975, 1985, 1995); Trepl (1987); Jax (2002); Schwarz (2003);
Kirchhoff 2007; Voigt 2009.
555 The Classical Holism-Reductionism Debate in Ecology
(1926) and Peus (1954) on the other, which (at least in a certain mode of
interpretation)24 express very clearly the essence of organicist holism and reductionist
individualism. An ideal type, after all, “is not a description of reality but it aims to
give unambiguous means of expression to such a description”.25 It “is formed by the
one-sided accentuation of one or more points of view and by the synthesis of a great
many diffuse, discrete, more or less present and occasionally absent concrete indi-
vidual phenomena, which are arranged according to those one-sidedly emphasized
viewpoints into a unified analytical construct”.26 The idea is – and this can only be
achieved through the method of constructing ideal types – to render classical
organicist holism and reductionist individualism comprehensible on the one hand,
and to formulate a foundation that makes it possible to describe and systematically
compare different concrete theories on the other.27
The Classical Organicist-Holistic Position
According to organicist holism, supra-organismic units, when described as “bio-
coenoses”, “communities” or “associations”, are communities that are determined
primarily by relationships of dependency between organisms.28 This dependency
refers to their development as well: Clements uses the term “reactions” to describe
the dependency of subsequent stages of succession on those preceding them. The
developing whole (see below) also depends heavily on the competitive activities of
the parts (Clements 1936, p. 143). In a stable end state (climax), however, the rela-
tionship between parts and whole consists essentially in each individual part being
brought forth by all the others, and therefore in a reciprocal bringing forth of the
24 For a detailed discussion of this mode of interpretation, in relation to Clements, see: Wolf, Judith
(1996): See also Eisel (1991).
25 “ist nicht die Darstellung des Wirklichen, aber er will der Darstellung eindeutige Ausdrucksmittel
verleihen” (Weber 1904, p. 234, emphasis in original).
26 “wird gewonnen durch einseitige Steigerung eines oder einiger Gesichtspunkte und durch
Zusammenschluß einer Fülle von diffus und diskret, hier mehr, dort weniger, stellenweise gar nicht,
vorhandenen Einzelerscheinungen, die sich jenen einseitig herausgehobenen Gesichtspunkten
fügen, zu einem in sich einheitlichen Gedankenbilde” (Weber 1904, p. 235, emphasis in original).
27 For the holism-reductionism controversy at the start of the twentieth century in ecology, see,
amongst others, McIntosh (1980); Tobey (1981); Worster (1985); Trepl (1987); Hagen (1992);
Golley (1993); and Anker (2001).
28 A biocoenosis and its habitat can also be seen together as comprising an organismic unit. “Jede
Lebensgemeinschaft bildet mit dem Lebensraum, den sie erfüllt, eine Einheit, und zwar eine in
sich oft so geschlossene Einheit, daß man sie gleichsam als einen Organismus höherer Ordnung
bezeichnen kann.” (Each community forms a unit with the habitat it fills; this unit is often so uni-
fied in itself that it can more or less be described as a higher-order organism.) (Thienemann and
Kieffer 1916, p. 485).
56 L. Trepl and A. Voigt
parts and the whole. Thus, individual organisms or even units made up of several
organisms, such as the functional groups of producers, consumers and decomposers,29
exercise certain functions within the community as a whole, much like organs in an
organism. These functions need to be carried out in order to preserve not just the
whole but also the organs themselves which carry out the functions. This kind of
community is a whole, which exists objectively, just like an individual organism, and
is “naturally” isolated and individual in space and time – if somewhat less clearly
than an individual organism.30 Moreover it is accorded the essential characteristics
of an organism – the “superorganism theory” is a prominent example of this. The
consequence of this for methodology is that the characteristics and behaviour of
individual organisms need to be examined according to the contribution they make
towards the functioning (adaptation) of the community as a whole.
Development of the Community as a Goal-Oriented Process
of Adaptation and Detachment
The organicist-holistic theory is a theory of development. This is usually not taken
into account, which means that the image of nature in holistic ecology is errone-
ously seen as a static one whenever – as occurs very often – an (allegedly outmoded)
29 “Nur die Pflanze kann, indem sie das Sonnenlicht als Energiequelle benutzt, aus anorganischen
Stoffen organische aufbauen und da das Tier sich nur von Organischem ernähren kann, so ist die
Tierwelt direkt oder indirekt fest mit der Vegetation verbunden. Wenn das Tier als Raubtier von
anderen tierischen Wesen lebt, so müssen die Beutetiere zur Lebensgemeinschaft des Raubtieres
gehören usw.” (Plants alone are able to form organic material out of inorganic material, using sun-
light as a source of energy, and since animals can only feed off organic material, the animal world is
firmly tied, directly or indirectly, to the vegetation. If an animal lives off other creatures as a predator,
then prey animals must be a part of the predator’s community etc.) (Thienemann 1944, pp. 7.)
30 “All diese Tiere und Pflanzen aber stehen nicht unvermittelt nebeneinander, sondern sind durch
die mannigfachsten Beziehungen aneinander gebunden; jede Stätte im Lebensraum hat so ihre
Lebensgemeinschaft oder Biocoenose.” (Yet all these animals and plants do not exist alongside
each other in an unmediated way; rather, they are tied to one another through many different
relationships. Thus every site in the habitat has its own community or biocoenosis.) (Thienemann
1944, p. 7). “So ist die Natur aufgegliedert in eine ganze Stufenfolge, eine Hierarchie ineinander
geschachtelter lebenserfüllter Räume, die nicht nur räumlich miteinander verbunden, sondern
auch voneinander durch den Kreislauf der Stoffe [...] abhängig sind. Jede Lebensstätte ist wie-
derum Glied einer größeren, bis hinauf zur ganzen Erde.” (Thus, nature is subdivided into a whole
layered succession, a hierarchy of interlinked spaces filled with life, which are not only connected
to each other spatially but are also dependent on one another due to the material cycle [...]. Every
living site is in turn part of a larger site, right up to the Earth as a whole.) (Thienemann 1944, pp. 8).
“Die Natur [...] ist, vom kleinsten Wiesenfleck angefangen bis zum ganzen Weltall, überall ein
geschlossener lebender Organismus, in dem jedes einzelne kleinste Glied auf jedes andere
abgestimmt ist; jede Veränderung eines Teils wirkt sich aus auf alle übrigen.” (From the smallest
spot of meadow to the whole universe, nature (...) is everywhere a unified living organism in which
every single tiny element is linked in with every other; any change in one part has an impact on
all the others.) (Thienemann 1944, pp. 35).
575 The Classical Holism-Reductionism Debate in Ecology
“ecological balance” is referred to (e.g. Pickett et al. 1992). This image of nature is
considered, in turn, to be one reason behind the existence of curiously static con-
ceptions in nature conservation (e.g. Reichholf 1993; Scherzinger 1996).
Just like an individual organism, the community develops according to intrinsic
rules of development through different “stages” to reach a state of maturity (cli-
max); it maintains itself in this state or else embarks anew on the process of devel-
opment: “As an organism the formation arises, grows, matures, and dies.” (Clements
1916, p. 3, our emphasis). The fact that this development is internally guided does
not mean that the community is autonomous, for it entails the latter differentiating
itself in accordance with external dictates while at the same time changing certain
of them. In the early phases of succession, those species that are adapted to the site
become established, that is, site conditions determine whether they will become
established. The communities made up of these species change the site, however.
They thus create conditions which are unfavourable for them but which will favour
species at later stages of succession. Having lost out in the competition, they are
replaced by the latter (cf. Clements 1916, 1936; Thienemann 1944). Unlike in
Darwinism, competition is not regarded from the perspective of individuals who
prove their worth in this “struggle for survival”, so that “progress” occurs in the
evolutionary line in terms of adaptation to particular environmental conditions;
instead, it is regarded in terms of the function it performs for the developmental unit
as a whole. The succession of different species at a certain site becomes an act of
self-sacrifice31 on the part of pioneer species for the benefit of “higher” stages and,
ultimately, the climax community.
From the perspective of the community, this process of replacement means that
it adapts as a whole to environmental conditions (“response”). This occurs above
all through a change in the composition of its species, prompted by competition,
and through internal differentiations, in particular an increase in the number of spe-
cies and in the number of interactions. At the same time, the community adapts the
locality to itself (“reactions”). “Each stage of succession plays some part in reducing
the extreme condition in which the sere began” (Clements 1916, p. 98). Since the
“stages” that manage to achieve this feat are superceded in this process, one can say
that the community adapts the locality to its own future constitution. In this way the
community becomes increasingly independent from specific site factors – from
those that exist initially in the sense that they no longer exist, and from those that
exist in later phases in the sense that they influence the organisms only indirectly,
through the community. Thus, development is simultaneously a process of adapta-
tion and detachment. The dependency of the whole on external factors decreases as
the dependency of the parts on one another and as the way they adjust to one
another increases. In contrast to the prior stages, the climax stage is such that it no
longer changes the external and internal conditions for the individual species in the
community. Instead, each individual organism “processes” the impacts of external
31 Der “Erhaltung des Ganzen wird, wenn nötig, auch das größte Teilglied geopfert.” (Even the largest
part will, if necessary, be sacrificed for the preservation of the whole.) (Thienemann 1944, p. 9).
58 L. Trepl and A. Voigt
factors through their interactions in such a way that the state of the whole remains
unchanged. Differentiation (that is, an increase in diversity), internal functional
dependency and mutual adjustment, and independence from external factors (in line
with the homeostasis of an individual organism) occur together in a necessary com-
mon context. The fact, that development can be characterised as internal means that
the community realizes that which was “embedded” in it from the start.
In the process of the site factors adapting to the developing community, these
factors are changed by the impact of the community in such a way that the original
diversity present at the site in question disappears (through the emergence of a
bioclimate, humus formation etc.). This then means that just one single climax
community will exist in each area governed by a specific climate, regardless of the
(original) small-scale differences at the site (what Clements called “monoclimax”,
and what Braun-Blanquet and Sukachev, for example, referred to in similar terms).
At the same time, the community optimizes its own adaptation to the conditions
presented by the regional climate in this development: in the climax state it exists
in a state of balance with this climate.32
Thus, this theory of development implies that the concept of the environment
divides into two categorically different concepts: (1) the many and varied small-
scale, temporally different site factors where the community lives (especially
edaphic and microclimatic factors), which in the course of succession relinquish
their influence on the development of the community and which the community
then adapts to itself; (2) the regional climate, over which it has no influence and to
which the community itself adapts in the course of its development. Adaptation is
complete in the climax state.
Thus, in organicist holism, succession is conceived of as being goal-oriented.
Even if we were not to interpret it in explicitly teleological terms, it is still teleo-
logical in the sense at least that it proceeds towards a final state conceived of as
being fixed from the start, because the regional climate – unlike the site factors – is
not modified by the community. This final state becomes established in spite of
different starting conditions (edaphic and micro-climatic differences, random
migration patterns). This theory is happy to accept that there are synecological units
which do not accord with the image of a highly integrated whole or with the
“intended” climax state. But by virtue of being a theory of development it manages
to conceptualise such units as partially developed phases or as deviations in the
“ontogenesis” of a community that is generally holistic, that is, one conceived of as
a superorganism.33
32 “[T]here is but one kind of climax, namely that controlled by climate” (Clements 1936, p. 128).
“Such a climax is permanent because of its entire harmony with a stable habitat. It will persist just
as long as the climate remains unchanged.” (Clements 1916, p. 99).
33 Clements sees successions that do not lead to a monoclimax community as deviations or as a
“subfinal stage of succession” (Clements 1936, p. 130). For example, there are very stable prior
stages (proclimax), subclimax or disclimax stages in which external natural factors or human
influences impede attainment of the climax stage (Clements 1916, 1936).
595 The Classical Holism-Reductionism Debate in Ecology
The Classical Individualist-Reductionist Position
For individualistic theories the individual organism is the fundamental unit which
alone is granted the status of “real”. The causal principle is regarded as being not only
permissible but indeed adequate for explaining the links between individuals and
between these and their abiotic environment. Thus, the notion of function in relation
to a whole that encompasses so many individuals is not what grounds this explana-
tion. In this respect, individualistic theories embody a form of biological mechani-
cism, even though the “fundamental units”, as emphasized above, are living
organisms. – Here again, we present an ideal typical reconstruction of this position.
We can take the following to be the starting point of the individualist position: “The
existence and success of each species depends solely on the realization and quality of
its own environment; it is left to fend for itself within that environment. There is nothing
above or beyond this that influences the animal and its life, ecologically speaking.”34
A community does not exist for the organisms.35 Other organisms are environmental
factors, as are abiotic factors; another living creature does not “appear” to an animal to
be such (Peus 1954, p. 300). From the perspective of the organism it is irrelevant, for
example, whether water is available in the form of a puddle (“abiotic factor”) or as a
component of a prey organism, as long as it is equally useful to it.36
This radical individualist position sees those units which ecologists call “bio-
coenoses”, “associations” etc. as “products of human imagination”37 (Peus 1954,
p. 300). Since they are fictitious they cannot be the object of scientific inquiry.
According to Peus ecology should restrict itself to autecology: “Biocoenology has
no grounding in reality as a science.”38 However, the logic of the individualistic
approach grants a certain heuristic usefulness to the concept of community. On the
one hand, communities must not be viewed as real entities, as they are in the
holistic-organicist view: not only are they not superorganisms, they are also not
“natural” units containing organisms that occur only in quite specific combinations
and that may be discovered and described – as individual species of organisms are –
by appropriate kinds of research. While it is true to say that individual organisms
34 “Jede Spezies ist in ihrer Existenz und in ihrem Gedeihen allein von der Verwirklichung und
Qualität ihrer eigenen Umwelt abhängig; sie ist darin auf sich allein gestellt. Darüber hinaus gibt
es nichts, was das Tier und sein Leben ökologisch gesehen beeinflußt.” (Peus 1954, p. 307).
35 This is not to say that relationships with other organisms may sometimes be very close, indeed
obligatory (obligatory predatory and mutualist relationships). What does not emerge, though, are
communities, i.e. units of a higher order in relation to which it would make sense, for example, to
say that the organisms were fulfilling certain functions for it. In the individualist view, it is only
in relation to individual organisms that it makes sense (heuristically) to say that a function is being
fulfilled for something.
36 This is why the individualist position is often linked to the “organism-centred approach”, which
attempts rigorously to adopt the perspective of the organism concerned in descriptions of environ-
mental factors (cf. Peus 1954; MacMahon et al. 1981; Jax 2002).
37 “Gebilde des menschlichen Vorstellungsvermögens” (Peus 1954, p. 300).
38 “Die Biozönologie als Wissenschaft hat keinen realen Grund” (Peus 1954, p. 300).
60 L. Trepl and A. Voigt
form groups with other organisms in a given area, the composition of these groups
will change depending on environmental factors and on the random nature of
migration patterns. On the other hand, however, it can be useful to give some of
them (e.g. those that appear more frequently under current environmental circum-
stances and conditions of migration) names (for example, names taken from the
system of plant sociology) in order to establish some point of reference amid the
many different combinations that occur. One just has to be aware that quite different
species combinations can also come about, and that the frequency of the species
groupings identified as “associations” etc. is due only to the coincidental fact that
the external circumstances required by these associations occur frequently – and
not to any internal rule of development which, even given very different conditions
at the start, gives rise time and again to specific combinations (such as that of the
“mature” state). In this view, there is no “structural uniformity of vegetation”, for
the simple reason that any given area is subject to annual variations (Gleason 1926,
p. 10). In addition, the issue of whether an area of vegetation is regarded as one
single association or as a mixture of several depends on the perspective taken by
the scientist concerned and on how the temporal and spatial boundaries are drawn
(Gleason 1926, pp. 10). When communities are isolated out by the scientist from
among the multitude of possible and real species combinations, other boundaries
and therefore other units (known as associations, for example) emerge, depending
on the focus of inquiry. A network of relationships will come to an end at different
points, depending on whether mutualistic or predatory relationships are selected for
study, so that the outcome is a different community. Viewed in this light, the scien-
tist is constructing communities.39
Changes to the Community Brought about by Changed
Environmental Conditions and the Randomness
of Species Migration
Views concerning the way in which “communities” (which may be nothing other
than groups of organisms occupying adjacent spaces) change through time differ
fundamentally in several ways within the individualistic-reductionist approach
39 The different individualist positions could be distinguished according to their starting assump-
tions. These may be (1) that communities can appear to be “naturally” demarcated – in other
words, individuals arrange themselves into certain groups according to, say, the requirements they
have of mutualistic partners. An objective boundary to the community is given at the point where
the (necessary) interactions finish. Alternatively (2) the boundaries may appear to be drawn by the
scientist according to his or her research interests. Within this, it is possible to distinguish
between a realistic and a nominalist variant. The realistic variant entails the view that there is a
(real) network of relationships that exists independently of an observer; this network has a specific
constitution, which is so complex, however, that we cannot (yet) recognize it and are forced, for
pragmatic reasons, to create “artificial” demarcations. The nominalist variant sees communities
fundamentally as being constructed by an observer (see also Chap. 27).
615 The Classical Holism-Reductionism Debate in Ecology
compared with the organicist-holistic approach where, as described above, changes
are seen as developments in the literal sense of the word:
1. “[Plants] can and do endure a considerable range in their environment” (Gleason
1926, p. 18). The occurrence of species in early phases of settlement is deter-
mined by the randomness of the arrival of diaspores and the existence of the
environmental conditions necessary for them to develop. Changes in the plants’
environment (site factors) are not primarily an outcome of the activity of organ-
isms,40 but occur above all by chance (e.g. climatic fluctuations, Gleason 1926,
p. 18). Therefore, the succession of different species combinations at any one
site is determined by the randomness of migration and by the randomness of
changing environmental conditions for the individual species. “The next vegeta-
tion will depend entirely on the nature of the immigration which takes place in
the particular period when environmental change reaches the critical stage”
(Gleason 1926, p. 21). Succession does not proceed, therefore, towards a par-
ticular end state (meaning: it brings to fruition that which was “inherent” in it
from the start); rather, it is random, it involves change but not development. If
immigration and environmental selection remain unchanged, then the commu-
nity attains stability; changes in one or both factors lead to a change in the com-
position of species. Climax – to the extent that this term makes sense in an
individualistic context – can mean no more than a phase in which no change
takes place for a certain period of time (Gleason 1926, p. 26). The further off in
the future it lies, the less the state of a community can be predicted, because one
cannot know enough about the environmental factors and migration events that
determine it. This contrasts with the organicist-holistic view, in which the ran-
dom and unpredictable nature of the initial conditions is emphasized, while the
end state is always the same: if we look far into the future, we can predict which
state will become established or, at any rate, which one ought “normally” to
become established, namely the climax state.
2. While it is a single community that develops in the organicist view, in the indi-
vidualist view it makes no sense to ask whether the phenomena observed consti-
tute a change in one or the succession of several communities – or rather, it can
be answered either one way or the other on account of the fundamentally arbi-
trary options for demarcation; it is simply that different combinations of species
follow on from one another in a more or less continuous rotation.
3. The adaptiveness of the individual organism is the starting point for the creation
of a community. Always and in every phase of succession, those organisms
become established that are adapted to the conditions which exist in that place.
It makes no sense to speak of the community adapting (with increasing success).
The role accorded to adaptation here differs radically from the one it plays in the
organicist approach. In the latter, adaptiveness (of the community to the regional
40 “The plant individual [...] is limited to a particular complex of environmental conditions, which
may be correlated with locality, or controlled, modified, or supplied by vegetation.” (Gleason
1926, p. 17).
62 L. Trepl and A. Voigt
climate) is the outcome of a development that leads to the climax community,
because the community increasingly gathers to itself those species which are
useful to it in the homeostatic maintenance of its organic balance.
4. In the individualistic approach, the concept of “law” has the meaning of a natural
law as commonly understood, in other words, a sentence containing the words
“whenever…, then…”, which always has to be known, along with the relevant
boundary conditions, whenever one wishes to explain or predict something.
Boundary conditions and (natural) law make it possible to explain why a particu-
lar change has occurred in the species combination at a site. In contrast to this,
the term “law” is used in a completely different sense in the organicist approach.
Here, changes in communities of organisms are described as “developments that
follow a set pattern” (“gesetzmäßige Entwicklungen”), meaning the same as
when the notion is applied to an organism, namely that in a “normal” or “typical”
case (in the normative, not statistical sense) certain conditions follow on from
one another. They are to be understood as a development of something which
was already “inherent” in the “undeveloped” state.
Can the Controversy Be Resolved?
We shall begin by discussing the hypothesis that there is no way empirically to resolv-
ing the holism-reductionism debate in ecology in favour of one or the other position.
We shall then discuss whether it might be possible to achieve a resolution of the issue
at the methodological level – perhaps one of the approaches uses inadmissible meth-
ods or methods that are poorly suited to the specific object of inquiry. In the course
of this discussion we will also deal with the problem of the admissibility of teleologi-
cal explanations of natural phenomena. The outcome of this discussion also suggests
that it is not possible to reach a clear decision on the issue. The controversy is there-
fore looked at from a third perspective: perhaps both approaches owe their existence
to external influences – “world views” – or are at least “inspired” by them, which
could mean that it is impossible to decide between them using scientific means.
The Controversy Cannot Be Resolved Empirically
All the evidence suggests that the dispute between organicist holism and reduction-
ist individualism cannot be resolved empirically, that is, by presenting facts that
would support the one and refute the other viewpoint. The assertions put forward
by each position can, as a rule, be explained just as well from the other side’s per-
spective. The holists maintain, for example, that a stable state is established at the
end of succession, and indeed they have been able to supply a certain amount of
empirical evidence to support this assertion. However, there are theories of the
635 The Classical Holism-Reductionism Debate in Ecology
individualist persuasion, which are equally able to explain these results as one possible
outcome of the course of successions (e.g. Horn 1976).41
To offer a second example, it has been argued from the individualist perspective
that empirical findings, if anything, fail to support the holistic assertion that suc-
cession follows certain laws, as suggested by the analogy with the development of
an organism (e.g. Drury and Ian Nisbet 1973). However, this argument basically
goes nowhere, insofar as holism doesn’t claim that the events formulated in the
law – such as an increase in the parameters of a community and in diversity during
the course of succession – can always or usually be observed in any given instance;
rather, it claims that those events “accord with” the healthy development of a
superorganism. Such developmental laws are thus of quite a different nature than
natural laws, such as the law of falling bodies, which have no exceptions. Indeed,
the claim that an increase in diversity follows a set pattern allows for any number
of “exceptions” to the rule, where the term “rule” does not describe what always or
normally happens (given specific parameters) but rather dictates what is supposed
to happen.
Thus an empirical resolution to the debate does not appear to be possible. It is
no surprise, therefore, that the dispute is often played out at a different level: the
other side is accused not so much of failing to supply sufficient empirical evidence
to support its claims as of using an inadmissible method.
Organicistic Holism, Individualist Reductionism
and the Problem of Teleology
It is the organicist approach in particular that is affected by the accusation of having
an inadequate methodology. While this approach is hard to refute at the empirical
level, it is certainly rather vulnerable methodologically. Indeed this is the main
objection that is levelled at every variant of holism. However, this objection is only
valid because (or rather, insofar as) its adherents are committed to the methodological
ideals of the modern natural sciences. Yet it is hard for them to avoid doing so42: It
can be seen as constitutive of modern thinking that each of the three ways of con-
templating nature – the aesthetic, the normative-evaluative and the scientific – are
valid independently of one another. This means that the scientific perspective ought
not to contain statements of the kind that characterise the normative-evaluative per-
spective. The constitutive role of the natural sciences is to provide causal explana-
tions and to avoid teleological ones. Values – and thus goals as well – can only be
41 On the reformulation of organicist-holistic arguments in the individualist context in general,
cf. Gnädinger 2002.
42 Explicit attempts of this kind have been made by Friederichs, for example (see above).
64 L. Trepl and A. Voigt
“attributed” to nature by us. Everything that is viewed from the scientific perspective
therefore appears to be a value-free object of theoretical knowledge.43
However, in biology, the science that deals with living nature, nature is often
spoken of in a way that seems to be diametrically opposed to the scientific method.
The concept of the living organism seems to include the fact that terms such as
benefit and harm, optimum, developmental goals etc. can be meaningfully applied
in the scientific context, which is not the case with non-living natural objects. The
organism (in modern societies, at least) is conceived of in such a way that each of
its parts owes its presence to the agency of all the other parts and exists for the sake
of the others and of the whole (Kant 1970, §§64–65). The parts are organs of the
whole. When one speaks of the function of the organs being to maintain the organ-
ism, one is making a teleological judgement. The term “self-maintenance”, when
used in relation to the organism, assumes that the state of life is the purpose and
desirable goal of the organism. Thus, it might appear that it is possible to attribute
a value to natural phenomena that is objective and is not related to human value
systems, but rather lies in the maintenance of the organism.
However, it is not possible in science to assert that things in nature happen
according to some purpose. The orientation towards purposes presupposes the idea
of a purpose which precedes a cause, i.e. action based on intention; yet we cannot
insinuate that nature has intentions. If we do, we are not looking at it scientifically.
Therefore, it is not only organicist holism in ecology but also the variant of holism
that concerns itself solely with individual organisms as an organic wholes – which
includes, in principle, the individualist position, as discussed above – that is vulner-
able to criticism from the perspective of radical mechanicism. To claim that natural
phenomena can be explained teleologically (and conceiving of them as the unity of
an organism is nothing else) would mean leaving behind the foundations on which
all the sciences rest.44
One possible objection to this radically mechanicist perspective is that its criti-
cism is unjustified if teleological explanations are intended heuristically and not as
objective explanations (cf. Kant 1970, §§69–78, see also above on methodological
holism). It is necessary to regard the organism as a whole and its parts as elements
that serve the function of maintaining it in order to have a point of reference for
biological research, even if the latter then has to explain its object in causal terms.
Indeed such explanations are indispensable in rendering the phenomenon of life
43 However, value freedom by no means signifies that as a general rule modern natural science is
not guided by interests, and hence by values. Its theories provide access to the “Wirklichkeit unter
dem leitenden Interesse an der möglichen informativen Sicherung und Erweiterung erfolgskon-
trollierten Handelns.” (reality under the guiding interest in assuring and extending instrumental
action in potentially informative ways.) (Habermas 1968, p. 157).
44 In the complex debate about this issue some positions claim that the problem of teleology is done
away with by evolutionary theory providing a causal explanation for the emergence of functional
characteristics (e.g. Mayr 1982), while others assert that teleological explanations can be recon-
structed as deductive-nomological ones (in Nagel’s sense), and that they should therefore defi-
nitely be accepted as scientific (cf. Looijen 2000).
655 The Classical Holism-Reductionism Debate in Ecology
“visible” at all (Cassirer 1921). Thus, if biology is to be a natural science, it must
use causal explanations. In order to justify the need for seeing biology as a special
kind of natural science, though, one is dependent on (heuristic) teleological judge-
ments (and therefore on methodological holism). This, in turn, can be held against
mechanicist reductionism: if the maintenance of the whole of an organism were not
conceptualised as constituting the purpose of the processes going on within it – even
if only in heuristic terms – then one would not be able to find a starting point for
physical-chemical explanations. In other words: physical-chemical explanations
would be irrelevant – they only acquire relevance when they contribute towards
explanations of the phenomena we call “life”.
The holistic approach in ecology, though, conceives not only of the individual
organism as an individual whole but the community and indeed “nature” as well.
This whole, for its part, consists of hierarchically interlocking individual wholes. It
is the function45 of the parts in the service of the whole (on which they in turn
depend) to work in such a way that everything supports everything else – just like
the relationship between organs and organism. Following Kant’s argument further,
however, it may be so that we are forced to use the notion of utility (even if only
heuristically) in the case of the (individual) organism, because the organism appears
to us to be “underdetermined” (that is, random) in its characteristics “according to
mechanical laws alone” (i.e. causally); but this is not the case with the community
of organisms (or with nature in general) (cf. Weil 2005). In order to explain why
the heart, for example, is constituted in a particular way, one needs to know what
function (what purpose) it fulfils in the organism. But in order to explain why cer-
tain plants grow in a community, one does not need to know their function in rela-
tion to other organisms, such as serving as food for animals – at most we need to
know the function other organisms have in relation to them. Even harder to com-
prehend is the call for explanations to be sought in the function they have in relation
to the whole of the community: while organs cannot live outside an organism,
organisms can, as a rule at least, live outside their respective community, namely in
other communities, or else in complete isolation. It is highly questionable, there-
fore, whether a teleological judgement relating to the whole of the community
makes sense, even if it is intended heuristically only. Ecological holism thus has a
bigger problem justifying its approach than does that of the physiologists, which
relates to the individual organism. The reason why it was – and still is – so wide-
spread in spite of this seems to call for explanation at a different level. This is what
we shall discuss in the following. What we seek to shed light on, though, is not only
the existence and even occasional dominance of organicist holism; rather, we shall
argue that the difficulty (or impossibility) of deciding between the two approaches –
the individualist approach is, as we have seen, just as vulnerable to critique at the
methodological level – can be better understood when one takes into account the
fact that both are constituted in part by external influences.
45 Etiological function as referred to by Wright (1973), cf. McLaughlin (2001).
66 L. Trepl and A. Voigt
Ecological Paradigms as Partially Constituted by Ideology
One way of bringing such external influences to bear theoretically is the hypothesis
that ecological holism is an effect of ideology – conservative ideology (cf. e.g. Eisel
1991). The major tenet of such ideologies is that “community” means an “organic”
community or a higher-order organism (Greiffenhagen 1971). The structural paral-
lels between conservative social theories and organicist ecological theories are
striking and have been widely discussed (e.g. Eisel 1991, 2002; Trepl 1993, 1997;
Körner 2000; Anker 2001; Schwarz 2003; Voigt 2009). The arguments fielded by
ecologists against organicist holism often seem to be grounded in the desire to keep
science pure from ideological contamination (e.g. Scherzinger 1995; cf. Körner
2000).
Looked at from the perspective of a theory of constitution, one important reason
why the organicist-holistic and individualist-reductionist positions exist in ecology
can be seen to lie in the fact that social relations exist which generate different
cultural ideas and make nature appear to be like this or like that. In the empirical
sciences, ideas that originated in culture are set up against one another and differ-
entiated as scientific theories, a move that generally goes unnoticed.
When the first theories about ecological units were developed in ecology, there
were two competing figures in philosophy, the social sciences and political theory
which constituted the framework for conceptualising the relationship between indi-
viduals and society. These can be categorised according to the two types of oppos-
ing world views that are constitutive of modernity, the one progress-oriented
(especially liberalism) and the other conservative. Even if these two basic figures
have been greatly changed, modernised and their components recombined in differ-
ent ways over time, both remain influential.
The liberal view (cf. e.g. Kühnl 1982; Arneson 1992) is that of a community of
autonomous individuals. It is based on the idea of an autonomous subject that has
liberated itself from the fetters of feudal society and of religion etc. so that now it
is responsible only for itself, pursuing its own advantage by means of general ratio-
nality and general technical knowledge. Rather than according with tradition and
nature, social development is related to individual emancipation and is therefore
open to progress. Liberalism is grounded in the idea that the world is shaped by the
struggle between individuals (Hobbes) who come together in communities because
this benefits them as individuals in the long term. In liberal theory society is seen
as the “sum” of individual people, who alone constitute reality. Society has no
“inner unity”; it is a superficial mechanical agglomeration of individuals. Society
is a system of interactions between individuals in which the interactions are geared
towards their usefulness for those individuals; it is a system in which interests are
reconciled in utilitarian fashion, premised on a “struggle for survival”.
This liberal view can be equated with the individualist approach in biology,
especially Darwinism and individualist ecological theories. Their structural con-
nections and, to some extent, the causal links in their emergence have been
675 The Classical Holism-Reductionism Debate in Ecology
described often.46 One aspect of their structural connections is that they both view
individuals’ needs as being the factor that forces them (individuals) to come
together in communities, that is, to establish relationships with others. In addition,
they both hold that social change has no ultimate goal: the direction it takes
depends upon chance factors, and if there is such a thing as development to a higher
level, then it does not involve a community coming closer to a pre-given goal but
rather an improvement from the perspective of individuals – of those who win
through in the competition for survival. Their respective ontological and epistemo-
logical views also match one another: it is a fundamental tenet of both individualist
ecology and liberalism and its related empiricist epistemologies that societies do
not exist in reality but are merely the outcome of the ordering activities of human
reason.
In contrast to this, the conservative view47 emphasizes the fact that the individual
is tied into a higher order – into an organic community (“Gemeinschaft”) rather than
just a community, or society, per se (“Gesellschaft”).48 The individual must accept
his or her predetermined ties to religion, nation, tradition, family etc., as it is only in
this constellation of relationships that the individual can develop his or her own
“special nature” (“Eigenart”). This is how each individual makes his or her own
contribution towards maintaining the God-given order of the whole (or that given by
nature or history) and this is how individuals contribute towards the development of this
whole, which is always organic and is always conceived of as “evolved”, rather than
as a construction. Each individual acts freely by recognizing and accepting the task
allotted to him or her in the given order. Freedom comes about, then, through the
recognition of ties. History is not an open process shaped by autonomous subjects
and determined by the forces of chance. Rather, it consists in the perfection of the
qualities inherent in “nations” (“Völker”) – their “character” – and in the perfection,
by these nations themselves, of the qualities inherent in the nature that exists in their
“Lebensraum” living space. This is the task allotted to nations as well as to individu-
als. “Culture” develops in the process of the nations adapting to the dictates of nature
in their “Lebensraum”, which simultaneously represents a process of breaking free
from the constraints of nature (cf. Kirchhoff 2005).
The structure of the counter-Enlightenment, conservative picture of community
displays a remarkably precise correspondence to the structure of the organicist-holistic
picture of ecological communities. Just as in political theories, the development
46 For Darwinismus cf. e.g. Engels (1886); Nordenskiöld (1928); Desmond and Moore (1991); for
the individualist approach in ecology e.g. Trepl (1994); Eisel (2002); Voigt (2009).
47 This is a reference to the basic conceptual figure of conservatism that arose in the course of the
counter-Enlightenment (cf. Eisel 1982). Later political movements grouped under the conservative
heading (e.g. “technocratic conservatism”) often deviate considerably from this (cf. Greiffenhagen
1971).
48 Cf. the distinction between “Gesellschaft“and “Gemeinschaft” in Tönnies (1887).
68 L. Trepl and A. Voigt
of the community represents the realization of that which is dictated to them at a
transcendental level (naturalised by Clements as the regional climate). This occurs
by means of creative individual effort, in a process of differentiation and the devel-
opment of particularity (“Eigenart”). In organicist ecological theory, too, the
development of a community is a process in which breaking free from the dictates
of nature (abiotic environmental conditions) occurs through adaptation to them.
Here, too, individuals are conceptualised as being not only physically dependent on
the whole per se (rather than on what they make for themselves out of the
“resources” available), but as being dependent on it also insofar as their existence
can only “make sense” in the context of the whole. This is because the meaning of
each person’s life lies in fulfilling the tasks given to them in relation to the whole
(within the context of the latter’s development towards the perfection of which it is
capable); it is a whole to which they owe their existence. Equally, the “purpose” of
individual organisms lies in exercising precisely those functions for the organic
community that are appropriate to the task of the self-maintenance of the commu-
nity as a whole and of its development towards the climax state.
Ecological paradigms can thus be read as political ideologies read back into the
workings of nature. Conversely, though, it is also the case that a particular ecologi-
cal paradigm entails particular political attitudes. Depending on how it is consid-
ered best to characterise ecological objects at a fundamental level, a certain set of
policies will appear to be meaningful and necessary. This applies not only to politi-
cal attitudes regarding the relationship between the individual and the society but
also, as we shall see, to the relationship between the society and nature. Having said
this, it would be rash to assume that a supporter of the organicist approach in ecol-
ogy, for example, must necessarily hold politically conservative views.49
Implications of Holism and Reduction
Whichever of the two views is taken as the basis for action, both entail far-reaching
practical consequences. This practical relevance may exist in relation to technology
(e.g. measures in agriculture and forestry, fishery, nature conservation) or in rela-
tion to “knowledge for orientation”, in which “nature” is viewed within a context
of values, either as being subject to evaluation or as a value-setting authority.
The Organicistic-Holist Approach in Nature Conservation
The organicistic-holistic position is a teleological one. At least, it is an obvious move
to interpret its empirical assertions in this way – that succession consists in a process
49 This would presuppose at the very least that for such a person an ideal community is oriented
towards an ideal image of nature. This is certainly usually the case, but is not necessarily so.
695 The Classical Holism-Reductionism Debate in Ecology
of differentiation leading to a climax state, and so on. In particular, it is hardly
plausible that a theory concerning the essence and the development of synecological
units could arise outside of a generally teleological conceptual structure, even if the
desire for scientific rigour may prompt attempts at causal reformulation.50 In a teleo-
logical interpretation of succession it is not simply that the community actually
changes in such a way that it moves towards a state of balance; rather, succession is
a development towards a goal.51 This climax state is not only a normal state in the
sense that it is generally reached; it is one that ought to be reached by the superorgan-
ism. Deviations from this are maldevelopments – similar developments in individual
organisms are judged no differently. The attributes associated with development – i.e.
internal differentiation and a consequent increase in diversity, functional integration
and stability in the sense of homeostatic independence from disrupting environmen-
tal impacts – are not merely facts in this view but are necessary attributes and the
standard for the development of the organic community to a higher development;
the climax state is the state of perfection (in relative terms, dependent on what is
possible for the community on the basis of its inherent “predispositions”). Thus,
value judgements are being (implicitly) expressed here. Initially, these are “values”
only from the perspective of the community (an end in itself). However, if “nature”
is conceptualised as an organic whole, just like the community, and if “the human
being” is seen as “a part of nature” – as is suggested by the world view generally
associated with that of ecological holism – then these values take on an ethical qual-
ity, because they are dictating a norm to humans.52
If the varieties of organisms in an organic community are all mutually dependent
on one another, then all the species are affected if one of them is removed, and the
obvious conclusion would be that the community as a whole has suffered harm.
However, every species (every organism) is judged on the basis of the function it
fulfils for the maintenance of the community; conversely, the whole is important
because it is indispensable for each individual species. Thus, the conclusion
reached in a circular but rigorous way from the premise that communities are
organisms is that “ecosystems” must be protected for the sake of the species and
species for the sake of ecosystems (on this, cf. especially biocentric-holistic
approaches to nature conservation of scholars such as Leopold 1949; Meyer-Abich
1984; Callicott 1989).
50 States of equilibrium at the end of a succession can in principle be understood in exactly the
same way as states of flow equilibrium found in the abiotic domain, that is, in purely causal terms.
However, in organicist theories they are understood as an organic balance (for this term, see Weil
1999) as in an organism. The question of whether one regards the end-stage explanations given
here as heuristic resources (see above) or whether one sees them as causes that are objectively at
work in nature is the deciding factor in whether or not theories about superorganisms (or theories
about organisms in general) can be regarded as scientifically admissible; whether they are correct
is another matter.
51 Cf. for early nature conservation e.g. Schoenichen (1942); Thienemann (1944).
52 On the different meanings of what is and can be called “value” and “judgement” around ecology
and nature protection, cf. e.g. Brenner (1996); Eser and Potthast (1997).
70 L. Trepl and A. Voigt
If change in a community constitutes a development much like that in an organism,
then what emerges from this is an indication of the kind of technology appropriate
to communities. The concept of the organism implies that the latter cannot be con-
structed but rather grows and develops. The organism can be nurtured – and it is
advisable to nurture it, if one wants to make “sustainable” use of it. Its development
can be encouraged, while deviations from the state it should be in require healing.
Such holistic natural objects may also be wholes consisting of nature and humans
(cultural landscapes).53 (cf. also Eisel 1980)
Arguments of this kind are widespread in both nature conservation and ecology,
as, for example, in current theories around the concept of “ecosystem health”.54
Indeed, it is hard to find any exceptions to what has become “common sense”
within the nature conservation and environmental movements. But one caveat
should be made at this point: the further away one goes from the sphere of commit-
ments and calls for environmental sensitivity and the closer one gets to the sphere
of administrative action, the more one finds elements that come from the opposite
(reductionist) direction – often in “bizarre and paradoxical rhetorical combina-
tions”55 (Hard 1994, p. 126).
The Individualist-Reductionist Approach
in Nature Conservation
The individualist approach has quite different implications due to its understanding
of the reality constituted by “community” (in the sense of “Gesellschaft”). What
should count as a community depends on the theories, definitions and questions
brought to bear by the scientist. Statements about a community being harmed or
destroyed are valid only in relation to the scientist’s definitions of what is to count
as a community in the first place. Similarly, the phenomenon of maldevelopment is
meaningful only in relation to these definitions. There is no state that could be
formulated as a point of reference on the basis of the requirements of the commu-
nity itself and in relation to which one might judge changes to be either positive or
negative. The community does not maintain itself against disturbances from outside
itself; rather, changes in the environmental conditions affecting individual species
bring about a rearrangement of species into a new group. There is no reason to
describe certain combinations as “intact” communities and others not. Since this
approach has no concept of a group of species as a community, as in the model of
an organism, it makes no sense to ascribe a “perspective” to a group of organisms,
53 Nothing should be added into the organic community, either. This results in a rejection of foreign
species (e.g. Disko 1996; for the debate about the role of foreign species in nature conservation,
see Eser and Potthast 1999, Körner 2000).
54 See. e.g. Rapport (1989, 1995); Costanza et al. (1992); Ferguson (1994); Rapport et al. (1998).
55 “bizarre und paradoxe rhetorische Mischungen”.
715 The Classical Holism-Reductionism Debate in Ecology
let alone one in which something may be “expedient” or “valuable” for the group
itself. Instead, in making such an ascription, one is necessarily relying on expecta-
tions directed at the system in question from the outside – the system may be useful
for particular purposes, but the purpose cannot lie in the system itself.56 It is not
possible, from the perspective of “communities”, to demand that they should not be
changed, but that they should be protected instead.57 This is because they do not
exist as living entities: one cannot meaningfully say that something is either good
or bad for them.
In the individualist perspective, species combinations are arbitrary as far as fur-
ther development is concerned; in contrast to organicism, they are not necessary life
phases in the development of a community towards the climax state.58 However,
they are not arbitrary in the sense that there are no causal laws governing the
arrangement of individual organisms into communities, because the abiotic envi-
ronment effects a selection and individuals act upon one another in a causal way
(competition, predator-prey relationships, etc.). If we knew the causal laws and the
parameters at work here, we could create specific species combinations, including
completely new ones. The technology appropriate to individualistically interpreted
communities is therefore one based on construction and not on nurture and
healing.59
Transcending the Dualism Between Holism and Reductionism?
Even if there is rarely any explicit mention of these classical positions in controver-
sies over the correct ecological theory and research practice, they can often be
reconstructed nonetheless as the axis around which the debate takes place. There
have been numerous attempts to adopt positions in between the two (e.g. Levins
and Lewontin 1980). Organicist holism is clearly discernible in those studies that
are commonly described under the rubrics of “ecosystem health” or “the Gaia
hypothesis” (e.g. Lovelock 1979). Apart from the latter, no one in today’s ecology
talks anymore about communities being superorganisms. Whenever this term is
used, it does not refer to the essence of communities in general, but only to certain
specific communities that are assumed to be units of selection (e.g. Wilson and
Sober 1989, Sober and Wilson 1994). Today’s ecology rarely contains such extreme
organicist-holist positions as those that were common during the first few decades of
the twentieth century. Changes have taken place since that time, including various
56 On the movements that have prompted such trains of thought in the theoretical development of
German environmental planning, see Eckebrecht (2002) among others.
57 The purpose we determine for them can also be, of course, to preserve a particular species.
58 This is why foreign species are not rejected in the corresponding conceptions of nature
conservation (e.g. Reichholf 1993).
59 The image of nature that comes to the fore here is hardly ever the individualistic one of a
“community of autonomous individuals”, but that of the machine, as we shall see below.
72 L. Trepl and A. Voigt
modernisations of the classical theories. For example, at the time when Clements’s
theory was highly influential in the USA, it was not so much its holistic character
in general that was criticised but rather the point that, even if different sets of cir-
cumstances existed in a climatic region, there could only be one single climax
community (“monoclimax”). The so-called polyclimax theory accorded local con-
ditions a much bigger role in the development of vegetation. Thus, it is possible for
different climax communities to come about in a single climax region, influenced,
for example, by small-scale varying edaphic factors (e.g. Tansley 1935, Whittaker
1953). The structure of the classical holistic approach can be seen more or less
clearly during the second half of the twentieth century, for example in the work of
E.P. Odum (1953, 1969), Margalef (1958), Patten (1978), Trojan (1984), Ulanowicz
(1997) and Jørgensen (2000), even if the organicist aspect usually remains secondary
to a more technical perspective in studies oriented towards systems theory (see
Chap. 27, as well as Chap. 15).
Individualist theories were initially paid scant attention.60 It was only during the
1950s that they assumed a more prominent role (e.g. Curtis and McIntosh 1950;
Egler 1951; Whittaker 1953), once certain approaches – sometimes described as
“population-oriented” – had been developed and began to be used in community
ecology. Ever since then, ecology has seen itself more within this tradition.61 Some
modern theories do exist that are very similar to the classical formulation of the
individualist concept (e.g. Hubbell 2001). However, rather than discussing the his-
tory of these approaches in any further detail, we intend in the following to look
instead at one important change to which holistic approaches were subject.
One factor that proved decisive for the further development of holism in ecology
was the coining of the ecosystem concept (Tansley 1935). Another equally impor-
tant factor – if not more so – was the attempt undertaken in other parts of biology
(especially in physiology) to accommodate the extreme reductionist and holistic
positions – encountered here predominantly in the form of an opposition between
mechanism and vitalism – within the framework of a systems theoretical concep-
tion of the organism, and later within General Systems Theory (Bertalanffy 1932,
1949, 1968)62. Both these factors laid the groundwork for the development in the
1940s and 1950s of a strand of research known as the ecosystem approach (see also
Chap. 15). Let it simply be noted here that ecosystem theories entail different sorts
of ideas about systems or, to put it another way, there is an apparent ambiguity
within the ecosystem concept, an ambiguity that is also found in General Systems
Theory with regard to the holism-reductionism issue (cf. Müller 1996).
Nowadays ecosystem theory comes in a bewildering array of variants.
Ecosystems are generally defined as open, self-regulating systems in a state of flow
60 For Anglo-American ecology, cf. McIntosh (1975, 1985, 1995); Simberloff (1980); for Europe,
see the fascinating debate around the essay by Peus (1954) in Schwerdtfeger et al. (1960/1961).
61 See also the debate recounted in Saarinen (1982).
62 For Bertalanffy’s early systems theory see Müller (1996); Schwarz (1996); Voigt (2001).
735 The Classical Holism-Reductionism Debate in Ecology
equilibrium, their defining characteristic being the uptake and output of matter and
energy (e.g. Lindeman 1942) and information (e.g. Margalef 1958).
Even if these ecosystem theories explicitly see themselves as holistic (e.g. Odum
1953; Jørgensen 2000; Jørgensen and Müller 2000) and even if they are holistic in
the sense that they take the whole of the system as given and see the parts as exist-
ing in a functional relationship to the latter, they are nonetheless reductionist in the
sense that they entail a considerable degree of abstraction and are resolutely scien-
tistic. Rather than focusing on communities (“Gesellschaften”) described as com-
binations of particular species, their object of study typically consists instead of
organisms or groups of organisms understood as “compartments” and viewed in the
same way as other components of the ecosystem, including abiotic ones, which
fulfil the same function within it.63
Those ecological theories from the second half of the twentieth century referred
to as holistic generally have this systems theoretical scientistic character (e.g.
Patten 1978; Jørgensen and Müller 2000) and it is this that distinguishes them fun-
damentally from the old organicist holism.64 There are, it is true, plenty of ecolo-
gists who use the concept of ecosystem and associate it with certain features of
communities described by the old organicist holism, such as that ecosystems are
real units which – like living beings – are self-enclosed, exist in relationship to their
environment, and develop like a unit towards individuality, so that in principle it is
possible to speak of their destruction in the same way we speak of the destruction
of an organism etc. There is also the view that the endogenous development of an
ecosystem leads to a predictable increase in biomass, diversity and stability in the
course of succession (particularly influential here: Odum 1969). To a certain extent
such theories involve reformulating the holistic-organicist concept of wholeness
within the framework of systems thinking (cf. McIntosh 1980). The systems con-
cept, which General Systems Theory takes as its starting point, proves attractive
insofar as it offers the possibility of conceptualising this (cf. Müller 1996).
However, the transition from organicism to ecosystem theory entailed a process of
scientization, the logic of which contains a tendency towards fundamentally changing
the focus of research and, with it, the kind of objects with which that research is
concerned (see Chap. 27).
The old organicist holism attempted to describe the entire range of organisms
occurring in a specific space as a higher-order organism, or at least to identify
“superorganisms” among the more or less close associations of individual organ-
isms. In other words, it sought at the synecological level those objects in nature
whose mode of functioning is aimed at self-maintenance. Ecosystem theory, by
contrast, demarcates certain systems, largely for instrumental reasons. These reasons
may be of a technical, practical nature or they may be theoretical. If they are the
latter, however, then they are also aimed at potential technical mastery: ecosystem
63 With regard to the laws that are formulated, it is often unclear, as in systems theory in general,
whether they are laws of nature – that is (usually physical) laws that relate to a particular domain of
the empirical world – or whether they are formal laws of the mathematical kind (cf. Müller 1996).
64 See the firm rejection of “ecosystems research” by such circles in Detering and Schwabe (1978).
74 L. Trepl and A. Voigt
theory regards ecosystems from the point of view of specific functions which the
system can fulfil for something outside of it, e.g. producing biomass, purifying
liquid wastes, stabilising the climate etc. (“ecosystem services”).65 In principle the
number of functions an ecosystem can fulfil is unlimited, as is the number of eco-
systems that can be formed by stating the functions from whose perspective they
have been defined. Thus, it is not the case that there are a certain number of (variet-
ies of) ecosystems (whose function lies in their self-maintenance), and which one
can discover and describe; rather, a fundamentally arbitrary number of (varieties of)
ecosystems are constructed (in theory or in reality), depending on particular inter-
ests, just like the communities (“Gesellschaften”) of the individualist approach.
Ecosystems are therefore artefacts,66 (super)organisms constitutively not. In mov-
ing from the concept of the superorganism to the concept of the ecosystem, the
purpose of ascribing value to certain states of the parts and the whole becomes a
slightly different one: the characteristics and behaviour of an ecosystem’s compo-
nents that are “good” for the ecosystem itself are those which make it possible for
them to fulfil the functions defined by us and by our interests. The structure of
(super)organisms is reflexive: the parts are self-generating, a process mediated via
the whole, and that is the function of both the parts and the whole. Superorganisms
decide for themselves, as it were, what is “good” for them. With ecosystems, how-
ever, we set an end point to what in principle is an endless cycle of functional utility,
and we do so at a point that seems interesting to us. By contrast, the chains of func-
tion in an organism reconnect to themselves, and this is why an organism is an end
in itself (cf. McLaughlin 2001). Despite the fact that the old organicist rhetoric
continues to be used in ecosystem theory (e.g. through terms such as self-regulation
and self-preservation as, for example, in Odum 1969), the meaning of those terms
has changed (even if they are often intended to convey their old meaning). One
refers thus to processes which, without any help from the outside, contribute
towards maintaining a state that is useful for the purpose defined by us.
These ecosystem theories can be understood and criticised as being both reduc-
tionistic and holistic. They are addressed towards the ecosystem as a whole but
reduce the diversity of its characteristics to a very few which, moreover, are amenable
to causal-mechanical analysis. The old problem of the relationship between parts and
wholes, or between individual and community, as it appears in the holism-reductionism
debate, is not addressed in these ecosystem theories, insofar as their focus is largely
on the material, energetic and possibly informational aspects of interactions in a
system, in which the difference between abiotic and biotic components is irrelevant:
the organisms actually involved in the interactions are not of interest (cf. Bergandi
1995; Voigt 2009). The idea abstracted from this is that the system’s elements (or the
biotic part of the system’s elements) are individual organisms of different species –
they are regarded merely in terms of their function (e.g. for the energy and material
65 Cf. e.g. the articles in Costanza et al. (1997); Daily (1997); de Groot et al. (2002); Farber et al.
(2002).
66 On this concept in relation to the concept of function in organisms, see McLaughlin (2001).
755 The Classical Holism-Reductionism Debate in Ecology
cycle of the system) which in turn is defined with regard to functions for some
utilitarian interest. This suggests that a different basic model has come to take the
place of the “organic community”, namely: the machine. cf. Voigt 2009.
Summary
Holism-reductionism controversies have long played a significant role in various
sciences, but especially so in philosophy and political ideologies. It is only against
this background that the holism-reductionism debate in ecology can be understood,
because here the issue is more than simply one of whether scientific theories of a
certain type describe certain natural phenomena correctly: it is a question of a con-
flict of ideas about the correct relationship between individual and community,
human beings and nature, progress and tradition. Taking this as our starting point,
we reconstruct the forms this controversy has assumed in ecology and which con-
ceptual structures it is based on; there is not an arbitrary range of these forms, but
rather only certain structurally feasible possibilities for conceptualising the rela-
tionship between parts and wholes.
Many different things are referred under the heading of holism and reduction-
ism. For example, it is possible to establish systematically those aspects of an
object of inquiry or of a scientist’s relationship to it to which holistic or reductionist
principles are being applied.
In ecology the whole is a community and the parts are individuals. The contrast
between reductionism and holism takes the form of an opposition between indi-
vidualism and organicism. These two approaches are presented here first as ideal
typical constructions in the classical form in which they emerged during the open-
ing decades of the twentieth century, using examples by way of illustration.
In organicist holism, “community” is conceived of specifically as an organic
community, or as a superorganism, that is, the relationship between parts and
wholes is conceptualised in analogy to the relationship between organ and organ-
ism. The key point is that the organicist-holistic theory is a theory of development.
The development of the community is a process – conceived of in teleological
terms – of adaptation to environmental conditions; at the same time it is a process
of breaking free from certain constraints. This corresponds closely to the structure
of conservative political philosophies: for the latter, too, society is an organic com-
munity, or a higher-order organism. Its development consists in the (active) perfecting
of the qualities inherent in “nations” (“Völker”) – their “character” – and in the
perfection of the qualities inherent in the nature that exists in their “Lebensraum”.
It is in adapting to the dictates of actual nature within their “Lebensraum” – which
simultaneously involves breaking free from the constraints of nature – that “culture”
develops. Rarely is any attention paid to the fact that holistic theory is also a theory
of development; the consequence is that holistic ecology’s image of nature is falsely
regarded as a static one, just like the image of society found in conservative political
philosophy.
76 L. Trepl and A. Voigt
As far as individualistic ecological theories are concerned, the fundamental unit
is the individual organism. They accord “reality” to it alone, not to the community.
It is the needs of the individuals, not the functional necessities of a community,
which force the individual organisms to establish relationships to others. Social
change is without a goal, its direction dependent on chance factors, and if there is
such a thing as development to a higher level, then this does not refer to a com-
munity coming closer its prescribed goal, but is instead an improvement from the
perspective of the individuals – those who win through in the competition.
Correspondingly, liberal political philosophies view society as a system of interac-
tions between individuals, where interaction results from its degree of usefulness
for the latter; it is also a system in which interests are reconciled in utilitarian fashion
on the premiss of a “struggle for survival”. History is an open-ended process shaped
by autonomous subjects and determined by the forces of chance.
Starting out from an approach based on a theory of constitution that allows us to
understand the two types of ecological theory as being “inspired” by political world
views, it is possible to understand better both the differences between the two types
and their immanent logic as well as the utterly divergent practical consequences
entailed by holistic ecological theories on the one hand and reductionist theories on
the other. “Practice” may be a reference to technology (e.g. “ecosystem manage-
ment”) or to “knowledge for orientation”, in which nature is viewed within a con-
text of values.
In conclusion, the dynamic that gives rise to both approaches and causes them
to change is described using the example of the transformation of the classical
holistic position. The old organicist holism sought at a synecological level those
objects in nature whose mode of functioning had the purpose of self-preservation.
In contrast to this, ecosystem theory – even when it explicitly calls itself holistic –
tends to demarcate certain systems in the interests of instrumental control.
Ecosystems are not superorganisms found in nature but rather artefacts. Through
this change, the purpose of attributing value terms to certain states of the parts and
the whole becomes a somewhat different one. A different basic model comes to
take the place of the “organic community”, namely: the machine.
References
Agazzi E (ed) (1991) The problem of reductionism in science. (Colloquium of the Swiss
Society of Logic and Philosophy of Science, Zürich, May 18–19, 1990. Zürich, Schweiz:
Schweizerische Gesellschaft für Logik und Philosophie der Wissenschaften). Kluwer
Academic, Dordrecht
Anker P (2001) Imperial ecology: environmental order in the British empire. Harvard University
Press, Cambridge, pp 1895–1945
Arneson RJ (ed) (1992) Liberalism 3 Vol. Schools of thought in politics 2 An Elgar reference
collection. - Hants: Edward Elgar, Aldershot (UK)
Ayala FJ (1974) Introduction. In: Ayala FJ, Dobzhansky T (eds) Studies in the philosophy of
biology: reductionism and related problems. Macmillan, London, pp 7–16
Ayala FJ, Dobzhansky T (eds) (1974) Studies in the philosophy of biology: reductionism and
related problems. Macmillan, London
775 The Classical Holism-Reductionism Debate in Ecology
Bertalanffy von L (1932) Theoretische Biologie, vol I, Allgemeine Theorie, Physikochemie,
Aufbau und Entwicklung des Organismus. Verlag Gebrüder Bornträger, Berlin, p 349
Bertalanffy von L (1949) Das biologische Weltbild, vol I, Die Stellung des Lebens in Natur und
Wissenschaft. Francke, Bern
Bertalanffy von L (1968) General system theory: foundations, development, applications.
Braziller, New York
Bergandi D (1995) “Reductionist holism”: an oxymoron or a philosophical chimaera of E. P.
Odum’s systems ecology? Ludus vitalis 3(5):145–180
Bergandi D, Blandin P (1998) Holism vs. reductionism: do ecosystem ecology and landscape
ecology clarify the debate? Acta Biotheor 46(3):185–206
Bews JW (1935) Human ecology. Oxford University Press, London
Bock GR, Goode JA (eds) (1998) The limits of reductionism in biology (Novartis Foundation sym-
posium 213, held at the Novartis Foundation, London, May 13–15 1997). Wiley, Chichester
Botkin DB (1990) Discordant harmonies: a new ecology for the twenty-first century. Oxford
University Press, New York
Braun-Blanquet J (1928) Pflanzensoziologie: Grundzüge der Vegetationskunde. J. Springer,
Berlin
Brenner A (1996) Ökologie-Ethik. Reclam, Leipzig
Bueno G (1990) Holismus. In: Sandkühler HJ (ed) Europäische Enzyklopädie zu Philosophie und
Wissenschaften. Felix Meiner Verlag, Hamburg, pp 552–559
Callicott JB (1989) In defense of the land ethic: essays in environmental philosophy. State
University of New York Press, Albany
Capra F (1982) The turning point: science, society, and the rising culture. Simon & Schuster,
New York
Cassirer E (1921) Kants Leben und Lehre. Verlag Bruno Cassirer, Berlin
Clements FE (1916) Plant succession: an analysis of the development of vegetation. Carnegie
Institution of Washington, Washington
Clements FE (1936) Nature and structure of the climax. In: The Journal of Ecology 24: pp 252–284
(– reprinted in: Allred B W & Edith S Clements (eds.) (1945) Dynamics of Vegetation: selec-
tions from the writings of Frederic E. Clements. – New York: The H. W. Wilson Company,
pp. 1–21
Clements FE, Shelford VE (1939) Bio-ecology. Wiley, New York
Costanza R, Norton BG, Haskell BD (eds) (1992) Ecosystem health: new goals for environmental
management. Island Press, Washington D.C.
Costanza R, d’Arge R, de Groot R, Farber S, Grasso M, Hannon B, Limburg K, Naeem S, O’Neill
RV, Paruelo J, Raskin RG, Sutton P, van den Belt M (1997) The value of the world’s ecosystem
services and natural capital. Nature 387(6230):253–260
Crick F (1966) Of molecules and men. (The John Danz lectures). University of Washington Press,
Seattle
Curtis JT, McIntosh RP (1950) The interrelations of certain analytic and synthetic phytosociologi-
cal characters. Ecology 31:434–455
Daily GC (ed) (1997) Nature’s services: societal dependence on natural ecosystems. Island Press,
Washington D.C.
Dawkins R (1976) The selfish gene. Oxford University Press, Oxford
de Groot RS, Wilson MA, Boumans RMJ (2002) A typology for the classification, description and
valuation of ecosystem functions, goods and services. Ecol Econ 41:393–408
Desmond A, Moore J (1991) Darwin. Michael Joseph, London, pp 21–807
Detering K, Schwabe GH (1978) System, Natur und Sprache. Scheidewege 8(1):104–132
Dilthey W (1883) Einleitung in die Geisteswissenschaften: Versuch einer Grundlegung für das
Studium der Gesellschaft und der Geschichte (1). Duncker & Humblot, Leipzig
Disko R (1996) Mehr Intoleranz gegen fremde Arten. Nationalpark 93(4):38–42
Drengson A, Inoue Y (eds) (1995) The deep ecology movement: an introductory anthology. North
Atlantic Books, Berkeley
Driesch H (1935) Die Maschine und der Organismus. In: Meyer-Abich A (ed) Bios 4. Barth, Leipzig
Drury WH, Ian Nisbet CT (1973) Succession. J Arnold Arboretum 54(3):331–368
78 L. Trepl and A. Voigt
Eckebrecht B (2002) Das Naturraumpotential. Zur Rekonstruktion einer geographischen
Fachprogrammatik in der Landschaftsplanung (Beiträge zur Kulturgeschichte der Natur 4. In:
Eisel U, Trepl L (eds) Freising: TU München, Lehrstuhl für Landschaftsökologie
Egler FE (1951) A commentary on American plant ecology based on the textbooks of 1947 – 1949.
Ecology 32:673–695
Ehrenfels von C (1890) Über Gestaltqualitäten. Vierteljahrsschrift für wissenschaftliche
Philosophie 14:249–292
Ehrenfels von C (1916) Kosmogonie. Eugen Diederichs, Jena
Eisel U (1980) Die Entwicklung der Anthropogeographie von einer “Raumwissenschaft” zur
Gesellschaftswissenschaft, vol 17, Urbs et Regio. Kasseler Schriften zur Geographie und
Planung, Kassel
Eisel U (1982) Die schöne Landschaft als kritische Utopie oder als konservatives Relikt. Soziale
Welt 38(2):157–168
Eisel U (1991) Warnung vor dem Leben. Gesellschaftstheorie als “Kritik der Politischen
Biologie”. In: Hassenpflug D (ed) Industrialismus und Ökoromantik: Geschichte und
Perspektiven der Ökologisierung. Deutscher Universitäts.-Verlag, Wiesbaden, pp 159–192
Eisel U (2002) Das Leben ist nicht einfach wegzudenken. In: Lotz A, Gnädinger J (eds) Wie
kommt die Ökologie zu ihren Gegenständen? Gegenstandskonstitution und Modellierung in
den ökologischen Wissenschaften. (Beiträge zur Jahrestagung des AK Theorie in der Ökologie.
– Theorie in der Ökologie 7). Peter Lang, Frankfurt a. M, pp 129–151
Engels, F (1886) Dialektik der Natur. In: Marx Karl and Friedrich Engels (1962) Werke, vol 20.
Dietz Verlag, Berlin, pp 305–570
Eser U, Potthast T (1997) Bewertungsproblem und Normbegriff in Ökologie und Naturschutz
aus wissenschaftsethischer Perspektive. Zeitschrift für Ökologie und Naturschutz
6:181–189
Eser U, Potthast T (1999) Naturschutzethik – Eine Einführung für die Praxis. Nomos-
Verlagsgesellschaft, Baden-Baden
Farber SC, Costanza R, Wilson MA (2002) Economic and ecological concepts for valuing ecosys-
tem services. Ecol Econ 4(3):375–392
Ferguson BK (1994) The concept of landscape health. J Environ Manage 40:129–137
Forbes SA (1887) In: The lake as a microcosm. Bulletin of the Scientific Association, Peoria,
pp. 77–87 (reprinted in 1925: Illinois Nat Hist Survey Bull. 15, 9: pp 537–550)
Foucault M (1969) L’ archéologie du savoir. Gallimard, Paris
Friederichs K (1927) Grundsätzliches über die Lebenseinheiten höherer Ordnung und den ökolo-
gischen Einheitsfaktor. Naturwissenschaften 15(7):153–157, 182–186
Friederichs K (1934) Vom Wesen der Ökologie. Sudhoffs Arch Gesch Med Naturwiss
27(3):277–285
Friederichs K (1937) Ökologie als Wissenschaft von der Natur oder biologische Raumforschung.
In: Bios 7. J. A. Barth, Leipzig
Friederichs K (1957) Der Gegenstand der Ökologie. Stud Gen 10(2):112–124, 10:3: 125–144
Gams H (1918) Prinzipienfragen der Vegetationsforschung Ein Beitrag zur Begriffsklärung und
Methodik der Biocoenologie. Vierteljahresschrift Naturforschende Gesellschaft Zürich
63:293–493
Gleason HA (1917) The structure and development of the plant association. Bull Torrey Botanical
Club 44:463–481
Gleason HA (1926) The individualistic concept of the plant association. Bull Torrey Botanical
Club 53:7–26
Gleason HA (1927) Further views on the succession-concept. Ecology 8(3):299–326
Gnädinger J (2002) Organismenzentrierte Rekonstruktion funktionaler Grenzen von synökologis-
chen Einheiten. In: Lotz A and Gnädinger J (eds) Wie kommt die Ökologie zu ihren
Gegenständen? Gegenstandskonstitution und Modellierung in den ökologischen Wissenschaften.
(Beiträge zur Jahrestagung des AK Theorie in der Ökologie. – Theorie in der Ökologie 7).
Peter Lang Verlag, Frankfurt a.M, pp 195–209
Goldstein K (1934) Der Aufbau des Organismus: Einführung in die Biologie unter besonderer
Berücksichtigung der Erfahrungen am kranken Menschen. M. Nijhoff, Haag
795 The Classical Holism-Reductionism Debate in Ecology
Golley FB (1993) A history of the ecosystem concept in ecology: more than the sum of the parts.
Yale University Press New Haven, New Haven
Greiffenhagen M (1971) Das Dilemma des Konservatismus in Deutschland. Piper, München
Grisebach A (1838) Über den Einfluß des Klimas auf die Begrenzung der natürlichen Floren.
Linnaea 12:159–200
Habermas J (1968) Erkenntnis und Interesse. In: Habermas, Jürgen: Technik und Wissenschaft als
“Ideologie”. Suhrkamp, Frankfurt a.M, pp 146–168
Hagen JB (1992) An entangled bank: the origins of ecosystem ecology. Rutgers University Press,
New Brunswick
Haldane JS (1931) The philosophical basis of biology (Donnellan lectures, University of Dublin
1930). Hodder and Stoughton, London
Hard G (1994) Die Natur, die Stadt und die Ökologie. Reflexionen über ”Stadtnatur“und
„Stadtökologie“. In: Ernste H (ed) Pathways to human ecology. Lang, Bern, pp 161–180
Harrington A (1996) Reenchanted science: holism in German culture from Wilhelm II to Hitler.
Princeton University Press, Princeton
Horn HS (1976) Succession. In: May RM (ed) Theoretical ecology: principles and applications.
Blackwell Scientific Pub. Ltd., Oxford, pp 187–204
Hubbell SP (2001) The unified neutral theory of biodiversity and biogeography. Princeton
University Press, Princeton
Hull DL, Ruse M (eds) (1998) The philosophy of biology. Oxford University Press, Oxford
Humboldt von A (1806) Ideen zu einer Physiognomik der Gewächse. Cotta, Tübingen
Jax K (1998) Holocoen and ecosystem on the origin and historical consequences of two concepts.
J Hist Biol 31:113–142
Jax K (2002) Die Einheiten der Ökologie: Analyse, Methodenentwicklung und Anwendung in
Ökologie und Naturschutz. Theorie in der Ökologie, 5th edn. Peter Lang, Frankfurt/M
Jørgensen SE (2000) A general outline of thermodynamic approaches to ecosystem theory. In:
Jørgensen SE, Felix M (eds) Handbook of ecosystem theories and management. Lewis
Publishers, London, pp 113–135
Jørgensen SE, Müller F (2000) Ecosystems as complex systems. In: Jørgensen SE, Felix M (eds)
Handbook of ecosystem theories and management. Lewis Publishers, London, pp 5–21
Kant I (1970) Kritik der Urteilskraft, edition 1995. Suhrkamp, Frankfurt/M
Keller DR, Golley FB (eds) (2000) The philosophy of ecology: from science to synthesis.
University of Georgia Press, Athens
Kirchhoff T (2005) Kultur als individuelles Mensch-Natur-Verhältnis. Herders Theorie kultureller
Eigenart und Vielfalt. In: Weingarten M (ed) Strukturierung von Raum und Landschaft.
Konzepte in Ökologie und der Theorie gesellschaftlicher Naturverhältnisse. Westfälisches
Dampfboot, Münster, pp 63–106
Kirchhoff T (2007) Systemauffassungen und biologische Theorien. Zur Herkunft von
Individualitätskonzeptionen und ihrer Bedeutung für die Theorie ökologischer Einheiten.
(= Beiträge zur Kulturgeschichte der Natur, Band 16). Freising
Köchy K (1997) Ganzheit und Wissenschaft: das historische Fallbeispiel der romantischen
Naturforschung, vol 180, Epistemata, Reihe Philosophie. Königshausen & Neumann, Würzburg
Köchy K (2003) Perspektiven des Organischen: Biophilosophie zwischen Natur- und
Wissenschaftsphilosophie. Schöningh, Paderborn
Köhler W (1920) Die physischen Gestalten in Ruhe und im stationären Zustand: eine naturphil-
osophische Untersuchung. Vieweg, Braunschweig
Körner S (2000) Das Heimische und das Fremde: Die Werte Vielfalt, Eigenart und Schönheit in
der konservativen und in der Liberal-progressiven Naturschutzauffassung. (Fremde Nähe
Beiträge zur interkulturellen Diskussion 14). LIT, Münster
Kuhn TS (1962) The structure of scientific revolutions. University of Chicago Press, Chicago
Kühnl R (1982) Das liberale Modell politischer Herrschaft. In: Abendroth W (ed) Einführung in
die politische Wissenschaft, 6th edn. Francke, München, pp 57–85
Langthaler R (1992) Organismus und Umwelt: die biologische Umweltlehre im Spiegel traditio-
neller Naturphilosophie, vol 34, Studien und Materialien zur Geschichte der Philosophie.
Georg Olms Verlag, Zürich
80 L. Trepl and A. Voigt
Lenoble F (1926) À propos des associations végétales. Bulletin de la Société Botanique de France
73:873–893
Leopold A (1949) A Sand County almanac: and Sketches here and there. Oxford University Press,
New York
Levins R, Lewontin RC (1980) Dialectics and reductionism in ecology. Synthese 43:47–78
Levins R, Lewontin RC (1994) Holism and reductionism in ecology. CNS 5(4):33–40
Lindeman RL (1942) The trophic-dynamic aspect of ecology. Ecology 23(4):399–418
Loeb J (1916) The organism as a whole: from a physicochemical viewpoint. Putnam’s Sons,
New York
Looijen RC (2000) Holism and reductionism in biology and ecology: the mutual dependence of
higher and lower level research programmes, vol 23, Episteme. Kluwer Academic Publisher,
Dordrecht
Lovelock JE (1979) Gaia: a new look at life on earth. Oxford University Press, Oxford
MacMahon JA, Schimpf DJ, Andersen DC, Smith KG, Bayn RLJ (1981) An organism-centered
approach to some community and ecosystem concepts. J Theor Biol 88(2):287–307
Margalef R (1958) Information theory in ecology. Gen Syst 3:36–71
Mayr E (1982) The growth of biological thought: diversity, evolution and inheritance. Belknap,
Cambridge
Mayr E (1988) The multiple meanings of teleological. In: Toward a new philosophy of biology: obser-
vations of an evolutionist. Belknap Press of Harvard University Press, Cambridge, pp 38–66
McIntosh RP (1975) H. A. Gleason. “Individualistic Ecologist” 1882–1975: his contributions to
ecological theory. Bull Torrey Botanical Club 102(5):253–273
McIntosh RP (1980) The background and some current problems of theoretical ecology. Synthese
43:195–255
McIntosh RP (1985) The background of ecology: concept and theory. Cambridge University
Press, Cambridge
McIntosh RP (1995) H. A. Gleason´s “individualistic concept” and theory of animal communties:
a continuing controversy. Biol Rev Camb Philos Soc 70:317–357
McLaughlin P (2001) What functions explain: functional explanation and self-reproducing sys-
tems. Cambridge University Press, Cambridge
Meyer-Abich A (1934) Ideen und Ideale der biologischen Erkenntnis. Beiträge zur Theorie und
Geschichte der biologischen Ideologien, vol 1, Bios. Barth, Leipzig
Meyer-Abich A (1948) Naturphilosophie auf neuen Wegen. Hippokrates, Stuttgart
Meyer-Abich KM (1984) Wege zum Frieden mit der Natur: praktische Naturphilosophie für die
Umweltpolitik. Hanser, München
Mittelstraß J (1995) Holismus. In: Mittelstraß J (ed) Enzyklopädie Philosophie und
Wissenschaftstheorie. Metzler, Stuttgart, pp 123–124
Mocek R (1974) Wilhelm Roux, Hans Driesch: Zur Geschichte der Entwicklungsphysiologie der
Tiere, Entwicklungsmechanik. Fischer, Jenas
Mocek R (1998) Die werdende Form: eine Geschichte der kausalen Morphologie, vol 3, Acta
biohistorica. Basilisken-Presse, Marburg
Müller K (1996) Allgemeine Systemtheorie. Geschichte, Methodologie und sozialwissenschaftliche
Heuristik eines Wissenschaftsprogramms, vol 164, Studien zur Sozialwissenschaft.
Westdeutscher Verlag, Opladen
Naess A (1973) The shallow and the deep, long-range ecology movement: a summary. Inquiry
16:95–100
Nagel E (1949) The meaning of reduction in the natural sciences. In: Stauffer RC (ed) Science and
Civilization. University of Wisconsin Press, Madison
Nagel E (1961) The structure of science: problems in the logic of scientific explanation. Harcourt,
Brace & World, New York
Nagel E (1979) Teleology revisited. In: Nagel E (ed) Teleology revisited and other essays in the
philosophy and history of science. Columbia University Press, New York, pp 275–316
Needham J (1932) Thoughts on the problem of biological organization. Scientia 52:84–92
815 The Classical Holism-Reductionism Debate in Ecology
Negri G (1928) Popolamento vegetale ed animale delle alte montagne: relazione illustrativa delle
proposte presentate dal Comitato Geografico Nazionale Italiano al Congresso internazionale
di Cambridge. Istituto geografico, Florenz militare
Nordenskiöld E (1928) The history of biology: a survey. (Originally issued as Biologins historia,
in three volumes, (1920–1924) Translated from the Swedish). Alfred A Knopf, New York
Odum EP (1953) Fundamentals of ecology. W. B. Saunders, Philadelphia
Odum EP (1969) The strategy of ecosystem development: an understanding of ecological succes-
sion provides a basis for resolving man’s conflict with nature. Science 164:262–270
Oppenheim P, Putnam HW (1958) Unity of science as a working hypothesis. In: Feigl H, Scriven
M, Grover M (eds) Concepts, theories and the mind-body problem. Minnesota studies in the
philosophy of science, 2nd edn. University of Minnesota Press, Minneapolis, pp 3–36
Patten BC (1978) Systems approach to the concept of environment. Ohio J Sci 78:206–222
Peus F (1954) Auflösung der Begriffe “Biotop” und “Biozönose”. Deutsche Entomologische
Zeitschrift 1:271–308
Phillips J (1934) Succession, development, the climax and the complex organism: an analysis of
concepts. Part I. J Ecol 22:554–571
Phillips J (1935) Succession, development, the climax and the complex organism: an analysis of
concepts Part II & III. J Ecol 23:210–246, 488–508
Pickett STA, Parker VT, Fiedler PL (1992) The new paradigm in ecology: implications for con-
servation biology above the species level. In: Fiedler PL, Jain SK (eds) Conservation biology.
The theory and practice of conservation, preservation and management. Chapman & Hall, New
York, pp 65–88
Portmann A (1948) Die Tiergestalt Studien über die Bedeutung der tierischen Erscheinung.
Reinhardt, Basel
Putnam H (1987) The many faces of realism. Open Court Publishing, La Salle
Ramensky LG (1926) Die Gesetzmäßigkeiten im Aufbau der Pflanzendecke. Botanisches
Centralblatt 7:453–455
Rapport DJ (1989) What constitutes ecosystem health? Perspect Biol Med 33(1):120–132
Rapport DJ (1995) Ecosystem health: more than a metaphor. Environ Values 4:287–309
Rapport DJ, Costanza R, McMichael AJ (1998) Assesing ecosystem health. Trends Ecol Evol
13(10):397–402
Reichholf JH (1993) Comeback der Biber: ökologische Überraschungen. Beck, München
Rosenberg A (1985) The structure of biological science. Cambridge University Press, Cambridge
Roux W (1895) Gesammelte Abhandlungen über Entwickelungsmechanik der Organismen, vol 2.
Wilhelm Engelmann, Leipzig
Ruse M (ed) (1973) Philosophy of biology. Hutchinson, London
Saarinen E (ed) (1982) Conceptual issues in ecology. Pallas paperback 23, Reidel
Scherzinger W (1995) Blickfang – Mitesser – Störenfriede. Nationalpark 88(3):52–56
Scherzinger W (1996) Naturschutz im Wald Qualitätsziele einer dynamischen Waldentwicklung.
Ulmer, Stuttgart
Schoenichen W (1942) Naturschutz als völkische und internationale Kulturaufgabe. Eine
Übersicht über die allgemeinen, die geologischen, botanischen, zoologischen und anthropolo-
gischen Probleme des heimatlichen wie des Weltnaturschutzes. Fischer, Jena
Schwarz AE (1996) Aus Gestalten werden Systeme: Frühe Systemtheorie in der Biologie. In:
Mathes K, Broder B, Klemens E (eds) Systemtheorie in der Ökologie. Beiträge zu einer
Tagung des Arbeitskreises “Theorie” in der Gesellschaft für Ökologie: Zur Entwicklung und
aktuellen Bedeutung der Systemtheorie in der Ökologie. ecomed, Landsberg, pp 35–45
Schwarz AE (2003) Wasserwüste Mikrokosmos Ökosystem. Eine Geschichte der ‘Eroberung’ des
Wasserraumes. Rombach, Freiburg im Breisgau
Schwerdtfeger F, Friederichs K, Kühnelt W, Illies JB, Schwenke W (1960/1961) Kolloquium über
Biozönose-Fragen. Z Angew Entomol 47:90–116
Simberloff DS (1980) A succession of paradigms in ecology: Essentialism to materialism and
probabilism. Synthese 43:3–39
82 L. Trepl and A. Voigt
Smuts JC (1926) Holism and evolution. Macmillan, New York
Sober E, Wilson DS (1994) A critical review of philosophical work on the units of selection problem.
Philos Sci 61:534–555
Stöckler M (1992) Reduktionismus. In: Ritter J, Gründer K (eds) Historisches Wörterbuch der
Philosophie 8. Schwabe, Basel, pp 378–383
Sukachev VN (1958) On the principles of genetic classification in biocenologie. Ecology
39:364–367
Tansley AG (1935) The use and abuse of vegetational concepts and terms. Ecology
16(3):284–307
Thienemann A (1941) Vom Wesen der Ökologie. Biologia Generalis 15:312–331
Thienemann A (1944) Der Mensch als Glied und Gestalter der Natur. Wilhelm Gronau, Jena
Thienemann A (1954) Ein drittes biozönotisches Grundprinzip. Arch Hydrobiol 49(3):421–422
Thienemann A, Kieffer JJ (1916) Schwedische Chironomiden. Arch Hydrobiol
2(Suppl):483–553
Tobey RC (1981) Saving the prairies the life cycle of the founding school of American plant ecol-
ogy, 1895–1955. University of Carlifonia Press, Berkley
Tönnies F (1887) Gemeinschaft und Gesellschaft: Abhandlung des Communismus und des
Socialismus als empirischer Culturformen. Fues, Leipzig
Trepl L (1987) Geschichte der Ökologie. Vom 17. Jahrhundert bis zur Gegenwart. Athenäum,
Frankfurt/M
Trepl L (1993) Was sich aus ökologischen Konzepten von “Gesellschaften” über die Gesellschaft
lernen läßt. Loccumer Protokolle 75(92):51–64
Trepl L (1994) Holism and reductionism in ecology: technical, political, and ideological implica-
tions. CNS 5(4):13–31
Trepl L (1997) Ökologie als konservative Naturwissenschaft. Von der schönen Landschaft zum
funktionierenden Ökosystem. In: Eisel U, Schultz H-D (eds) Eographisches Denken, vol 65,
Urbs et Regio, pp 467–492
Trojan P (1984) Ecosystem homeostasis. Dr. W. Junk Publishers, the Hague
Troll W (1941) Gestalt und Urbild: Gesammelte Aufsätze zu Grundfragen der organischen
Morphologie. Akademische Verlagsgesellschaft, Leipzig
Ulanowicz RE (1997) Ecology, the ascendent perspective. Columbia University Press, New York
Uexküll von J (1920) Staatsbiologie: Anatomie, Physiologie. Pathologie des Staates. Berlin,
Paetel
Voigt A (2001) Ludwig von Bertalanffy: Die Verwissenschaftlichung des Holismus in der
Systemtheorie. Verhandlungen zur Geschichte und Theorie der Biologie 7:33–47
Voigt A (2009) Die Konstruktion der Natur. Ökologische Theorien und politische Philosophien
der Vergesellschaftung. Franz Steiner, Stuttgart
Weber M (1904) Die “Objektivität” sozialwissenschaftlicher und sozialpolitischer Erkenntnis. In:
Weber M (1988) Gesammelte Aufsätze zur Wissenschaftslehre. 7. (ed) Tübingen: Mohr,
Siebeck, pp 146–214
Weil A (1999) Über den Begriff des Gleichgewichts in der Ökologie Ein Typisierungsvorschlag.
In: Trepl L (ed) Gleichgewicht – Funktion der Biodiversität. Landschaftsentwicklung und
Umweltforschung, vol 112. TU Berlin, Berlin, pp 7–97
Weil A (2005) Das Modell “Organismus” in der Ökologie: Möglichkeiten und Grenzen der Beschreibung
synökologischer Einheiten, vol 11, Theorie in der Ökologie. Peter Lang, Frankfurt/M
Wertheimer M (1912) Experimentelle Studien über das Sehen von Bewegung. Barth, Leipzig
Whittaker RH (1953) A consideration of climax theory: the climax as a population and pattern.
Ecol Monogr 23:41–78
Wilson DS, Sober E (1989) Reviving the superorganism. J Theor Biol 136:337–356
Wolf J (1996) Die Monoklimaxtheorie: Das biologische Konzept vom Superorganismus als
Entwicklungstheorie von Individualität und Eigenart durch Expansion. In: Naturalismus.
Projektbericht in zwei Bänden. S. 231–308. Bd. I. TU Berlin, Fachbereich 7
835 The Classical Holism-Reductionism Debate in Ecology
Wolf KL, Troll W (1940) Goethes morphologischer Auftrag. Versuch einer naturwissenschaftli-
chen Morphologie. Akademische Verlagsgesellschaft, Leipzig
Worster D (1985) Nature’s economy: a history of ecological ideas. Studies in environment and
history. Cambridge University Press, Cambridge
Wright L (1973) Functions. In: Conceptual issues in evolutionary biology, 2 edn. MIT Press,
Cambridge, pp 27–49 (reprinted in: Sober E (ed) (1994))
Part III
About the Inner Structure
of Ecology – Some Theses
87
Introduction
My thesis is that, although ecologists have not named the concept as such, they are
always dealing with unruly complexity, that is, with ongoing change in the structure
of situations that have built up over time from heterogeneous components and are
embedded or situated within wider dynamics. Ecology tends to suppress such com-
plexity by mimicking the physical sciences in constructing – materially and con-
ceptually – well-bounded systems, which have clearly defined boundaries, coherent
internal dynamics, and simply mediated relations with their external context.
Ecologists can envisage themselves positioned outside the systems and seek gener-
alizations and principles that afford a natural or economical reduction of complex-
ity. If researchers want, in contrast, to discipline unruly complexity without
suppressing it, they need to recognize that control and generalization are difficult
and no privileged standpoint exists; that ongoing assessment is needed, and this
requires engagement in the changing situations. The inner structure of ecology
rests, therefore, on the tension between unruliness and attempts to discipline it.
This article, which builds on Taylor and Haila (2001), reviews the recent history
of ecological theory with a view to highlighting the challenges of conceptualizing
heterogeneity, embeddedness, and ongoing restructuring. The subject matter of this
review is not well-bounded; at places I point well beyond the terrain of ecology
proper. Indeed, the HOEK project of illuminating ecological concepts by position-
ing them in relation to the socio-historical context in which they are produced and
deployed invites us to consider embeddedness of other kinds: ecological situations
within socio-environmental processes; natural science within the systematic study
of social change; conceptual work within scientific practice; and interpretation of
science within engagement in scientific and social change. All these interconnected
realms pose analogous challenges of dealing with unruly complexities (Taylor
2005). Although the unruliness-system tension can be seen in other disciplines,
Chapter 6
Conceptualizing the Heterogeneity,
Embeddedness, and Ongoing Restructuring
That Make Ecological Complexity ‘Unruly’
Peter Taylor
P. Taylor ()
University of Massachusetts Boston, USA
e-mail: peter.taylor@umb.edu
A. Schwarz and K. Jax (eds.), Ecology Revisited: Reflecting on Concepts, Advancing Science,
DOI 10.1007/978-90-481-9744-6_6, © Springer Science+Business Media B.V. 2011
88 P. Taylor
such as evolutionary biology, epidemiology, and developmental psychology, ecology
provides a fruitful entry point for exploring this epistemic type.
A Brief History of Recent Ecological Theory
Let me acknowledge at the outset that my conceptual map is centred in the United
States and needs to be balanced by reference to other HOEK entries (see also
Chaps. 7 and 8). During the 1960s and 1970s many U.S. ecologists sought theories
of ecological structure and function that would be general and not dependent on
historical particularities (Kingsland 1995, pp. 176–205). Systems ecologists,
through extensions of thermodynamics and information theory to open biological
systems, sought to explain complexity in terms of the nutrient, energy, and informa-
tion flows within entire ecosystems (Taylor 2005). Community ecologists made
theoretical propositions, often expressed as mathematical models, which focused
on the regulation of population sizes and distributions through competition for
limiting resources and other interactions. The two schools mapped broadly onto a
series of conceptual-methodological contrasts: function and process vs. structure
and demography; properties of wholes vs. explaining parts and building up from
there; field measurements vs. mathematical modeling (Hagen 1989; see Taylor
1992 for a more complex map of commonalities and distinctions).
The status of model building was somewhat ambiguous: were models idealized
representations of ecological reality (see, e.g., the “perfect crystals” of May 1973)
or heuristic devices to generate further theoretical questions (Levins 1966)?
Moreover, by the early 1980s ecologists of a particularistic bent were questioning
many of community ecology’s models, rejecting them when their fit to data was no
better than alternative “null” hypotheses or “random” models (Strong et al. 1984).
Scepticism about the possibility of general ecological theory became widely
expressed. As Simberloff (1982) argued: Many factors operate in nature and in any
particular case at least some of them will be significant. A model cannot capture the
relevant factors and still have general application. Instead, ecologists should inten-
sively investigate the natural history of particular situations and test specific
hypotheses about these situations experimentally. They may be guided by knowl-
edge about similar cases and may add to that knowledge, but they should not expect
their results to be extrapolated readily to many other situations.
To some extent the particularistic current of the 1980s had been prefigured in
plant ecology’s shift from predictable stages of succession to shifting associations
of individual species determined by their particular life histories and environmental
requirements (McIntosh 1985; Taylor 1992 - or even earlier in the ecologically-rich
third chapter of Darwin’s 1859 “On the Origin of Species”). In other ways, how-
ever, vegetation ecology’s long tradition of descriptive studies was leading to a
different understanding of the difficulty of theorizing about ecological processes.
Multivariate statistical techniques (or pattern analysis) could be used to cluster
ecological sites into distinct communities (classification) or position them
along continuous axes (ordination). The patterns exposed could then be used to
896 Conceptualizing the Heterogeneity, Embeddedness, and Ongoing Restructuring
generate hypotheses about causal factors or underlying environmental gradients. By
the 1980s, however, vegetation ecologists (especially in Australia) had shown that
the results of pattern analyses were sensitive to the models underlying the technique
used and the sampling sites from the space of environmental possibilities. Popular
techniques, such as principal components analysis and detrended correspondence
analysis, when tested on simulated data, did not recover well the simulated environ-
mental gradients. Techniques that reduce this model-dependence also tend to pro-
duce degenerate patterns (Faith et al. 1987; Minchin 1987). The Catch-22 is that
one needs to know a lot about the causal factors behind the data in order to design
efficient and distortion-free multivariate techniques that would expose those factors
(Austin 1987). Inferring process from pattern is a problem that I remark on later,
after further discussion of particularism versus modeling and theory-building.
The scepticism about theory and a one-sided emphasis on hypothesis testing that
gained attention in the 1980s have been resisted from several angles: Observation and
experiment can contribute to the generation of theory in ways other than through
crucial hypothesis tests. Indeed, observations constructed for testing of a specific
hypothesis may not be useful for thinking about anything beyond the local configura-
tion observed. Theory generation draws on the many other faces of data: initial cate-
gory-generating generalizations from observations, comparisons, analytic
redescriptions (Haila 1988; see also Chap. 7 for an analytic description of ecological
concepts and theories). Moreover, exploration of verbal and mathematical models has
a valuable role in generating new concepts, framings, questions, and hypotheses.
An example of exploration of models, which is relevant to unruly complexity,
concerns investigation of how complexity of communities is related to their persis-
tence or stability. Originally, ecological theory implied that ecological complexity
was able to persist because of the enhanced stability of complex ecological systems.
However, mathematical analysis during the 1970s and 1980s showed that complexity
works strongly against stability unless the complexity is nearly decomposable, i.e.,
consists of loosely linked subsystems. Subsequently, a “landscape” view arose,
which holds that a community may persist in a landscape of interconnected patches
even though the community is transient in each of the patches (DeAngelis and
Waterhouse 1987). Meta-population theory, an actively explored variant, examines
the persistence not of communities, but of populations (or phoretic associations of
communities on carrier species) in such a landscape (Hastings and Harrison 1994).
Another variant of the landscape view emerges from construction of model systems
by addition and elimination of populations. This exploration shows that complexity
can persist—at levels far greater than found in decomposable systems – even when
any particular system is transient. Investigations of ecological complexity should
incorporate continuing turnover of species, not only analysis of the stability and
structure of the current configuration (Nee 1990; Taylor 2005). (Notice, however,
that the investigation of assembly rules in community ecology – see Weiher and
Keddy (1999) – tends to conflate pattern and process. In the absence of information
about historical trajectories, assembly rules are better thought of as patterns of co-
occurrence that are statistically significantly different from patterns that are pro-
duced by randomly sampling – “assembling” – species from the appropriately
delimited species pool; Kelt and Brown 1999.)
90 P. Taylor
By reintroducing historical contingency, transient or non-equilibrium situations,
and embeddedness in larger contexts, exploratory modeling of the construction and
turnover of systems, among other factors, contributes to undermining the aspirations
of earlier decades for identifying general principles about systems and communities
(Kingsland 1995, pp. 213–251). Since the 1980s ecologists in general have become
increasingly aware that situations may vary according to historical trajectories that
have led to them; that particularities of place and connections among places matter;
that time and place is a matter of scales that differ among co-occurring species; that
variation among individuals can qualitatively alter the ecological process; that this
variation is a result of ongoing differentiation occurring within populations – which
are specifically located and inter-connected – and that interactions among the spe-
cies under study can be artifacts of the indirect effects of other “hidden” species.
In patch dynamic studies, for example, the scale and frequency of disturbances
that create open “patches” is now emphasized as much as species interactions in the
periods between disturbances (Pickett and White 1985). Studies of succession and
of the immigration and extinction dynamics for habitat patches pay attention to the
particulars of species dispersal and the habitat being colonized, and how these deter-
mine successful colonization for different species (Gray et al. 1987). On a larger
scale such a shift in focus is supported by biogeographic comparisons that show that
continental floras and faunas are not necessarily in equilibrium with the extant envi-
ronmental conditions (Haila and Järvinen 1990). From a different angle, models that
distinguish among individual organisms (in their characteristics and spatial location)
have been shown to generate certain observed ecological patterns, such as patterns
of change in size distribution of individuals in a population over time, where large
scale, aggregated models have not (Huston et al. 1988; Lomnicki 1988). And, the
effects mediated through the dynamics of populations not immediately in focus, or,
more generally, through “hidden variables,” upset the methodology of observing
the direct interactions among populations and confound many principles, such as the
competitive exclusion principle, derived on that basis (Wootton 1994).
Hidden variables and indirect effects have potentially profound consequences
for conceptualizing ecology. Consider the strategy of scientific simplification in
which models refer only to a few populations, even though those populations are
embedded in naturally variable and complex ecological situations. Unless ecolo-
gists know that the full community has been specified, their “simple” models –
including “null” models – are primarily redescriptions of the particular observations
that do not provide, through their fit or lack of fit, sure or general insight about
actual ecological relationships. It should be noted that progress in the physical sci-
ences depends greatly on controlled experiments, in which systems are isolated
from their context. Yet this model of science is not appropriate for understanding
organisms embedded in a dynamic ecological context and responding to resources
and hazards that are unevenly distributed across place and time (Taylor 2005).
Embeddedness and the confounding effects of hidden variables should prod
theoreticians to scrutinize the analogies and conceptual borrowings drawn from
work on well-bounded systems. Similarly, the heterogeneity of units in ecology and
their disparate temporal and spatial scales of activity limits the relevance of
916 Conceptualizing the Heterogeneity, Embeddedness, and Ongoing Restructuring
complexity theory in which iterations of simple rules over time and space lead to
complex behaviors for which parallels in real-life are suggested (Waldrop 1992).
Long-standing physical and chemical theories in which macro-regularities arise
statistically from large numbers of similar entities also seem problematic for
linking patterns of ecological complexity and corresponding processes.
Extending thermodynamics to open “systems of man and nature” was, indeed, a
central motivation for H. T. Odum’s pioneering contributions to systems ecology
(Taylor 2005). Although the search for theoretical principles became less important
in subsequent systems ecology (during the 1960s and 1970s, especially during the
International Biological Programme), it reemerged in the 1980s in accounts of eco-
logical complexity as a hierarchy of systems embedded within larger systems with
complementary processes and patterns at each level or scale. One hope of hierarchy
theory was that, if the right measure were found for extracting patterns from data, a
natural reduction of complexity might be achieved (Allen and Starr 1982).
To some extent the problem of inferring process from pattern can be overcome
through the use of analysis of variance and related statistical techniques on data
from replicated, multi-factorial field experiments (Underwood 1997). Strictly
speaking, however, such results are local, that is, contingent on the configuration of
other factors held experimentally or statistically constant for the experiment
(Lewontin 1974). Localization poses few problems when ecological engineering
affords control over conditions and isolates the system from any surrounding
dynamics. But these are special cases.
Moving from Systems to Intersecting Processes
For some ecologists the growing emphasis since the 1980s on situated, scale-
crossing processes means that ecology needs to be reconceived as an “historical”
science (Schluter and Ricklefs 1993). Like epidemiologists, paleontologists, and
historians, ecologists face the challenge of historical explanation. That is, they have
to assemble a composite of past conditions sufficient for the subsequent outcomes to
have followed and not some other, while, at the same time, they must not obscure the
provisionality of such accounts in the face of competition from other plausibly suf-
ficient accounts (Miller 1991). This “composite of past conditions” would include
considerable historical and geographical contingency (such as which organisms sur-
vived in pockets when Mt. St. Helens erupted, Franklin and MacMahon 2000) and
the evolutionary particularity or “individuality” of species (Sterelny and Griffiths
1999, pp. 253 ff.). Yet historicity need not eliminate ideas about regularities or struc-
turedness of ecological patterns and processes. To say that ecological structure has a
history could be to say that it changes in structure and is subject to contingent, spa-
tially located events, while at the same time the structure constrains and facilitates the
living activity that constitute any ecological phenomenon in its particular place.
Whether or not the label “historical” is used, a key challenge for conceptualizing
ecology is to allow for particularity and contingency intersecting with structure, and
92 P. Taylor
for that structure to change, be internally differentiated, and, because of overlapping
scales of different species’ activities, have problematic boundaries. Systems that are
well bounded or have simple relations with their external context, when they are
encountered, could be viewed not as simple situations, but as special cases whose
existence requires explanation.
It is here that conceptual clarification may benefit from viewing ecological situ-
ations as special cases of intersecting social-environmental processes, which would
mean giving less weight to cases in which human disturbance is minimal or con-
stant, and allowing the natural science of ecology to be informed by debates in
social science. In particular, there is a “political ecological” current of anthropo-
logical and geographical research that focuses on situated, scale-crossing socio-
environmental processes. This research analyzes environmental problems in terms
of intersecting economic, social and ecological processes, which operate across
various spatial and temporal scales and are mutually implicated in the production
of any outcome and in their own ongoing transformation (Taylor and García-
Barrios 1995; Peet and Watts 1996). Accounts of soil erosion or collapse of fish
stocks, for example, may tie together the local and regional ecological characteris-
tics, local institutions of production and associated agro- or aqua-ecologies, the
social differentiation in a given community and its social psychology of norms and
reciprocal expectations, and national and international political economic changes
(Little 1987; García-Barrios and García-Barrios 1990; Taylor 2005).
Researchers who analyze “intersecting processes” have not articulated a mature
conceptual or methodological framework, but explanations that preserve heteroge-
neity of causes and complexity of their inter-linkages warrant much more attention
from philosophers and ecologists. Conceptual work in this area would require,
among other things, attention to researchers’ practice and engagement with the
complexity studied (Haila and Levins 1992; Goldman et al. 2011). Moreover,
intersecting processes accounts expose multiple points of potential engagement –
each one partial in the sense of being insufficient to overcome the focal problem,
and thus needing to be inter-linked within the ongoing intersecting processes
(Taylor 2005).
Partiality is pertinent even when researchers do not focus on socio-environmental
dynamics, but confine themselves to natural ecology. The exploratory use of models,
mentioned earlier, retains support, in part, because of an unstated implication that,
if the different exploratory models could be combined, they would yield an under-
standing of ecological phenomena that could not be achieved through the construc-
tion of all-encompassing systems models. For example, the idea that there is a limit
to the similarity of co-existing species might be combined with the ideas that spatial
heterogeneity or an intermediate level of disturbance promote diversity, and so on.
But how? The means of weaving together or synthesizing necessarily partial mod-
els, or heuristics, has yet to be articulated. On the reasonable assumption that few
ecologists can juggle more than a few heuristics, new approaches to conceptual
work need to be developed that bring different types of ecologists into sustained
interaction (Lee 1993; Walters 1997; Wondolleck and Yaffee 2000; Resilience
Alliance, http://www.resalliance.org (04.06.2007)).
936 Conceptualizing the Heterogeneity, Embeddedness, and Ongoing Restructuring
Social Interactions in the Production of Ecological Knowledge
Self-consciousness about social interactions involved in producing knowledge
lay behind Levins’ (1966) strategy of modeling, which distinguished his perspec-
tive from contemporaries in mathematical ecology. Levins has been concerned
with the vitality of the modeling process – with a never-ending process of distur
bing the provisional validity of models (Taylor 2000). His interest in the circum-
stances under which theoretical principles might be overthrown – circumstances
that are not always apparent to scientists – has led him to consider the social
conditions in which knowledge is produced (Haila and Levins 1992). For exam-
ple, under a research and development system geared to firms making profits,
pesticides have been favored over biological control of pests (Levins and
Lewontin 1985, pp. 238–241).
Socio-historical contextualization should, as a matter of logical consistency,
apply to any HOEK-style interpretation of ecological concepts. Indeed, I locate the
origins of my interest in unruly complexity at the turbullent intersection of ecology-
as-science and ecology-as-social-action during the 1970s. During this period ecol-
ogy-as-social-action involved a serious critique of the scientific enterprise. This
challenged researchers not only to attend to environmental concerns, but also to
shape scientific practices and products self-consciously so as to contribute to trans-
forming the dominant structure of social and environmental relations. This context
led me to engage with the complexities of environmental, scientific, and social
change together, as part of one project. The intersections of these three kinds of
change conditioned me to emphasize, both conceptually and in practice, problem-
atic boundaries, heterogeneity of ecological and social agents, and continuing
process over competed product (Taylor 2005).
It is relevant to describe here one aspect of the subsequent conceptual evolution
up to my current status as a HOEK contributor. I first attempted to interpret science
in its socio-historical context during the 1980s when I examined H. T. Odum’s
efforts to reduce the complexity of social and ecological relations to a single cur-
rency – energy – whose flows could be adjusted or redesigned. I associated this
with the post-war climate of “technocratic optimism” and proposed Odum had
found in nature a special role for systems engineers, such as himself, working in the
service of society (Taylor 2005). Although I had shown that the social embedded-
ness of science can have systematic effects on the content of scientific knowledge,
the scientist in me wanted to develop ways to bring such interpretations to bear
productively on subsequent research. To provide insights about how that might be
achieved, a finer-grained analysis than the broad historical interpretation of Odum
seemed to be ca yze shorter-term projects of socio-environmental assessment likely
to be governed by more complex and contested pragmatics (Bocking 1997). During
this interpretive work, my image of scientists working in a social context evolved
from “social-personal-scientific correlations” into one where scientists have to deal
with diverse considerations in practice. This allowed me to emphasize a range of
different points at which researchers, interpreters of science as well as scientists,
94 P. Taylor
could engage differently in scientific practice and try to modify its outcomes.
Whether any specific modifications are do-able depends on the position and
resources of the specific researchers – myself included – as they enter into negotia-
tions with other relevant social agents. The ability of researchers in practice to
make knowledge is distributed beyond their persons, not concentrated mentally
inside them; it depends on intersecting processes (Taylor 2005).
How do we get to know ecological complexity? The answers depend on paying
more attention to who “we” are, to the associations different people make as they
position themselves – in practical as well as conceptual terms - in relation to life’s
complex, changing, and rarely well-bounded ecological context. The inner structure
and process of ecology is surely a matter of heterogeneity, embeddedness, and
ongoing restructuring.
References
Allen TFH, Starr TB (1982) Hierarchy. perspectives for ecological theory. University of Chicago
Press, Chicago
Austin MP (1987) Models for the analysis of species’ response to environmental gradients.
Vegetatio 69:35–45
Bocking S (1997) Ecologists and environmental politics. A history of contemporary ecology. Yale
University Press, New Haven
Darwin C (1964/1859) On the origin of species. Harvard University Press, Cambridge
DeAngelis DL, Waterhouse JC (1987) Equilibrium and non-equilibrium concepts in ecological
models. Ecol Monogr 57:1–21
Faith DP, Minchin PR, Belbin L (1987) Compositional dissimilarity as a robust measure of eco-
logical distance. Vegetatio 69:57–68
Franklin JF, MacMahon JA (2000) Messages from a mountain. Science 288:1183–1185
García-Barrios R, García-Barrios L (1990) Environmental and technological degradation in peas-
ant agriculture. A consequence of development in Mexico. World Dev 18(11):1569–1585
Goldman MJ, Nadasdy P, Turner MD (eds) (2011) Knowing Nature: Conversations between
Political Ecology and Science Studies, University of Chicago Press, Chicago
Gray AJ, Crawley MJ, Edwards PJ (eds) (1987). Colonization, succession and stability. 26th
Symposium of the British ecological society. Blackwell, Oxford
Hagen JB (1989) Research perspectives and the anomalous status of modern ecology. Biol Philos
4:433–455
Haila Y (1988) The multiple faces of ecological theory and data. Oikos 53:408–411
Haila Y, Järvinen O (1990) Northern conifer forests and their bird species assemblages. In: Keast
A (ed) Biogeography and ecology of forest bird communities. SPB Academic, the Hague,
pp 61–85
Haila Y, Levins R (1992) Humanity and nature. Ecology, science and society. Pluto, London
Hastings A, Harrison S (1994) Metapopulation dynamics and genetics. Annu Rev Ecol Syst
25:167–188
Huston M, DeAngelis D, Post W (1988) From individuals to ecosystems. A new approach to
ecological theory. Bioscience 38:682–691
Kelt DA, Brown JH (1999) Community structure and assembly rules. Confronting conceptual and
statistical issues with data on desert rodents. In: Weiher E, Keddy P (eds) Ecological assembly
rules, perspectives, advances, retreats. Cambridge University Press, Cambridge, pp 75–107
Kingsland S (1995) Modeling nature: Episodes in the history of population biology. University of
Chicago Press, Chicago
956 Conceptualizing the Heterogeneity, Embeddedness, and Ongoing Restructuring
Lee K (1993) Compass and gyroscope. Integrating science and politics for the environment.
Island Press, Washington DC
Levins R (1966) The strategy of model building in population biology. Am Sci 54:421–431
Levins R, Lewontin R (1985) The dialectical biologist. Harvard University Press, Cambridge
Lewontin RC (1974) The analysis of variance and the analysis of causes. Am J Hum Genet
26:400–411
Little P (1987) Land use conflicts in the agricultural/pastoral borderlands. The case of Kenya. In:
Little P, Horowitz M, Nyerges A (eds) Lands at risk in the third world. Local-level perspec-
tives. Westview Press, Boulder, pp 195–212
Lomnicki A (1988) Population ecology of individuals. Princeton University Press, Princeton
May RM (1973) Stability and complexity in model ecosystems. Princeton University Press,
Princeton
McIntosh RP (1985) The background of ecology. Concept and theory. Cambridge University
Press, Cambridge
Miller RW (1991) Fact and method in the social sciences. In: Boyd R, Gasper P, Trout JD (eds)
The philosophy of science. MIT Press, Cambridge, pp 743–762
Minchin PR (1987) An evaluation of the relative robustness of techniques for ecological ordination.
Vegetatio 69:89–107
Nee S (1990) Community construction. Trends Ecol Evol 2:337–343
Peet R, Watts M (eds) (1996) Liberation ecologies. Environment, development, social movements.
Routledge, London
Pickett STA, White PS (eds) (1985) The ecology of natural disturbance and patch dynamics.
Academic, Orlando
Schluter D, Ricklefs R (1993) Species diversity. An introduction to the problem. In: Ricklefs R,
Schluter D (eds) Species diversity in ecological communities. University of Chicago Press,
Chicago, pp 1–10
Simberloff D (1982) The status of competition theory in ecology. Ann Zool Fenn 19:241–253
Sterelny K, Griffiths P (1999) Adaptation, ecology, and the environment. Sex and death. An intro-
duction to the philosophy of biology. University of Chicago Press, Chicago
Strong DR, Simberloff D, Abele LG, Thistle AB (eds) (1984) Ecological communities, conceptual
issues and the evidence. Princeton University Press, Princeton
Taylor PJ (1992) Community. In: Keller EF, Lloyd E (eds) Keywords in evolutionary biology.
Harvard University Press, Cambridge, pp 52–60
Taylor PJ (2000) Socio-ecological webs and sites of sociality. Levins’ strategy of model building
revisited. Biol Philos 15(2):197–210
Taylor PJ (2005) Unruly complexity: Ecology, interpretation, engagement. University of Chicago
Press, Chicago
Taylor PJ, García-Barrios R (1995) The social analysis of ecological change. From systems to
intersecting processes. Soc Sc Info 34(1):5–30
Taylor PJ, Haila Y (2001) Situatedness and problematic boundaries. Conceptualizing life’s com-
plex ecological context. Biol Philos 16(4):521–532
Underwood AJ (1997) Experiments in ecology. Their logical design and interpretation using
analysis of variance. Cambridge University Press, Cambridge
Waldrop MM (1992) Complexity. The emerging science at the edge of order and chaos. Simon
and Schuster, New York
Walters C (1997) Challenges in adaptive management of riparian and coastal ecosystems. Conserv
Ecol 1(2):1, http://www.consecol.org/vol1/iss2/art1
Weiher E, Keddy P (eds) (1999) Ecological assembly rules. Perspectives, advances, retreats.
Cambridge University Press, Cambridge
Wondolleck JM, Yaffee SL (2000) Making collaboration work. Lessons from innovation in natural
resource management. Island Press, Washington DC
Wootton JT (1994) The nature and consequences of indirect effects in ecological communities.
Annu Rev Ecol Syst 25:443–466
97
Introduction
Ecology is a pluralistic science (McIntosh 1985, 1987; Cherrett 1989; Dodson
1998). Pluralism can be explained by the following hypotheses:
It is a result of recent diversification. Applications in environmental sciences lead
to differentiation within an accepted theoretical framework (centrifugal model).
It is a result of a long historic development. Incorporation of disparate branches of
sciences were accompanied by incomplete theory reduction and unification
(attractor model).
It is a result of a complete lack of coherent theory formation. Disagreement on
aims, methods and accepted theories prevails (anarchy model).
I assume in this context that the third possibility holds. Any overview of ecology
requires a delimitation towards nonecology. Here I define the current set of accepted
facts, theories and laws in ecology according to the contents of the textbooks Begon
et al. (1986) and Krebs (1995). This circumscription will help us to trace back
ecological approaches before the rise of self-conscious ecology. In accordance with
Begon et al. I define ecology as the science that deals with the description, explana-
tion and prediction of individuals, populations, and communities in space and time.
Variables of interest are distribution, amount (biomass), number (abundance, diver-
sity) and composition (similarity) of biotic entities (Peters 1991; Krebs 1995).
First I will focus on the internal structure of ecology with special reference to its
historical roots. I present a short outline of my personal approach towards history
of science. I then give an overview of the main phases of the development of ecol-
ogy including an investigation of the importance of benchmark papers and books
since 1920. In the second part I investigate the fields in which current ecologists
work. Those are defined by systematic groups, habitat types, observational levels,
spatial scales, research programmes, schools and traditions, key concepts, and
applications. Criteria such as the distinction between reductionistic vs. holistic
Chapter 7
A Few Theses Regarding the Inner
Structure of Ecology
Gerhard Wiegleb
G. Wiegleb (*)
Chair General Ecology, Brandenburg University of Technology, Cottbus, Germany
e-mail: wiegleb@tu-cottbus.de
A. Schwarz and K. Jax (eds.), Ecology Revisited: Reflecting on Concepts, Advancing Science,
DOI 10.1007/978-90-481-9744-6_7, © Springer Science+Business Media B.V. 2011
98 G. Wiegleb
ecology (Wilson 1988), between organismic vs. individualistic ecology (Simberloff
1980; Trepl 1987), competing methodological approaches (experimental, comparative,
exploratory, simulation, Grime 1979), or competing aims (explanation, prediction,
description; Wiegleb 1989, Peters 1991) are used for distinguishing research
programmes, schools, or preferences for key concepts.
A Short History of Ecology
General Aspects
History of ecology is regarded as the history of ecological ideas (May and Seger
1986; Trepl 1987). The theory of Kuhn (1976), assuming scientific revolutions and
subsequent change of paradigms (= complete replacement of valid ideas), is not
applicable to ecology. According to Trepl (1987) ecology never developed any
paradigm comparable to physics, chemistry, or molecular biology. Simberloff
(1980) distinguished two paradigms in ecology, the organismic and the individual-
istic one. However, both sets of ideas cannot be regarded as paradigms in a Kuhnian
sense. Or we have to assume the coexistence of two different, noncommunicating
sciences. Neither can the “paradigms” of Regier and Rapport (1978) be seriously
considered as Kuhnian paradigms.
I am inclined to follow Lakatos (1978) in assuming coexistence of competing
research programmes and stepwise replacement of degenerative approaches by more
promising ones. In ecology, research programmes were not deliberately formulated
but developed as a casual consequence of benchmark papers or textbooks. Thus they
are only recognizable in retrospective. According to Toulmin (1978) scientific con-
tinuity can be recognized on the levels of theory, content, sociology, and psychology.
Questions such as the following deserve an answer: Which were the factors of inno-
vation in the past which produced preliminary variants of ideas? What was the con-
text of discovery? Which were the factors of selection fixing final variants of ideas?
What was the context of agreement? Which factors prevented the break-through of
ideas? Additionally, the approach of Mayr (1984) contributes significantly to gaining
insight into historical processes. He describes the growth of ideas in individual per-
sons, asking: Which scientists were exceptional, what did they think? How did
researchers proceed to achieve their results? Which were the individual influences
on their work on a biographical level? He thoroughly investigates the psychological
and sociological hindrances of agreement upon theories and facts.
In this brief historical account the interplay between the development of ecol-
ogy, the progress in philosophy of science, and the societal context cannot be
analysed in detail. Progress in epistemology and philosophy of science between
Paracelsus, Bacon and Descartes (empirism vs. rationalism) and Leibniz, Hume
and Kant played an important role in the early separation of biology from natural
history (until 1790). Later inner-biological developments and the accumulation
of facts played a great role (until 1940). More recently, external factors relating
ecological ideas to the development of social groups, institutions, politics, and
997 A Few Theses Regarding the Inner Structure of Ecology
technology have become more important. To date those chapters of the history of
ecology are mostly unwritten (except for Küppers et al. 1978; Golley 1993;
Anker 2001).
Early Phases of Ecology Development
Here I distinguish between seven phases of ecology, which have left their traces in
recent ecological theory and practise. Besides autochthonous development, external
scientific traditions were incorporated in the course of time.
Phase 1. During ancient to medieval times (600 BC to AD 1300) science was part
of philosophy. Not much genuine ecological knowledge was accumulated. It is
restricted to medicinal botany (from Dioscorides to Hildegard von Bingen 1150) and
population studies (since 1200, L. Pisano; P. de Crescenzi; Egerton 1973, 1983).
Phase 2. During early modern times (from 15th to middle of eighteenth century)
several traditions developed which paved the way for the achievements of Linnaeus.
The most prominent are:
Around 1,520 botany was revived by so-called “herbalists”. Several nicely illus- –
trated herbal books appeared between 1,520 and 1,540. Authors such as O. Brunsfels,
H. Bock, L. Fuchs, A. Mattioli, R. Dodonaeus, Ch. de L’Écluse, and V. Cordus
partly included pedological observations and descriptions of the growth place of
plants (Mägdefrau 1992).
Encyclopedic natural history started in the sixteenth century with its most –
famous representatives K. Gesner, U. Aldrovandi, P. Belon and G. Rondelet
(Jahn et al. 1985).
Early phytogeography of the seventeenth century borrowed both from herbalism –
(Fuchs) and natural history sources (Gesner). Important representatives are
B. Varenius and J.P. de Tournefort (Egerton 1977).
Harmonistic viewpoints of nature date back to ancient times, e.g. Pythagoras –
(Egerton 1973). They were revived at the end of the seventeenth century by
J. Ray, who in his “Natural history of animals” advanced an explicit application
of the “Balance of Nature” concept to natural history (Jansen 1972; Egerton
1973). W. Derham, a student of Ray’s, developed “Physicotheology” into a com-
prehensive system which was discussed as a serious theory until 1850, in par-
ticular in England, later seeking confrontation with Darwinism. As a part of
religious and environmentalist ways of thinking it is still recognizable in the
current discussion on environmental protection.
Realistic paintings of plants and insects (still life) influenced natural history dur- –
ing the seventeenth and early eighteenth century. Based on earlier Dutch exam-
ples A.S.M. Merian and J. Rösel von Rosenhof edited coloured books showing
animals feeding on plants. Linnaeus used such material for describing species
(Jahn et al. 1985).
Starting around 1,580 the work on population development by F. Botero, D. Peteau, –
Th. Browne, J. Graunt, M. Hale and D. Dodart culminated 1798 in T.R. Malthus’
100 G. Wiegleb
treatment “Essay on the Principle of Population”. He postulated geometric
growth of human population leading to famine and disease (“misery and vice”).
The mathematical-theoretical population ecology was important for the origin of
evolution theory, but it remained an independent line of thought until the beginning
of the twentieth century (R. Pearl, A. Lotka, V. Volterra). Only around 1,970 an
amalgamation to the mainstream of ecology occurred (Egerton 1976, 1977).
Phase 3. The scientific revolution during the age of enlightenment and the origin of
science in a modern sense (around 1,750) lead to the protoecological work of
C. von Linnaeus and some predecessors in the framework of natural history.
Important predecessors of the early eighteenth century were A. van Leeuwenhoek, –
studying dynamics of microbes, R. Bradley, founding production biology, and
R.A.F. de Reaumur, founding animal ecophysiology and ecophysiology of
plants (Abbot 1983; Egerton 1969, 1977).
Linnaeus contributed to various disciplines of ecology, e.g. floristics, vegetation –
geography (description of Skandinavian vegetation in terms of altitudinal stages,
zonal stages and environmental gradients), mire science (description of mire
types), lake science (description of lake types according to the vegetation, specula-
tions on nutrient content of the lakes), indicator plants (formulation of the indica-
tor principle), succession of plant communities (in bog waters), food chains
(relationships between animals and their host plants, observations on the function
of carnivores), experimental ecology (planting of plants of foreign countries in
Sweden imported by his students), dispersal ecology (speculations on creation
centres in disjunct species), and phenology (Bremekamp 1952; DuRietz 1957).
The general importance of Linnaeus might be seen in his distinction between –
religion and science. He treated the “Balance of Nature” concept as a scientific
theory rather than as a proof of divine wisdom (Egerton 1973; Querner 1980).
Further achievements of Linnaeus were the foundation of biological systematics
and the delimitation of the science from other cultural enterprises such as medi-
cine, pharmacy, agriculture, and cooking. In his work, Linnaeus combined
observation, theory formation and experiment.
Phase 4. Development of “self-conscious ecology” in the nineteenth century was
based on the influence of two main lines of scientific development (Worster 1977;
Trepl 1987; see also Chap. 4).
The development of biogeography, in particular plant geography is based on the –
seminal work of Alexander von Humboldt (1806), later followed by De Candolle
(père et fils), Schouw, Meyen, Grisebach, Kerner von Marilaun, Drude, and
Schimper in plant geography, and Latreille and Wallace in animal geography.
(Nelson 1978; Jahn et al. 1985). Animal geography was trailing behind and
reached the mainstream only around 1920 (Shelford 1913).
Evolutionary biology as developed by J.B. de Lamarck and C. Darwin between –
1800 and 1860 paved the way for ecological thought. While Lamarck’s theory
was physiologically oriented, Darwin presented a theory in which external factors
were responsible for organism change (Stauffer 1960; Vorzimmer 1965; Egerton
1968; Wuketits 1995).
1017 A Few Theses Regarding the Inner Structure of Ecology
Theoretical considerations by E. Haeckel and practical contributions by –
K. Möbius (Reise 1980) and V. Hensen (Lussenhop 1974) between 1866 and
1878 lead to the development of the first ecological research programme by
Warming (1895); (Goodland 1975).
Phase 5. Soon after the publication of Warming’s book (between the edition of the
English translation in 1909 and 1925), ecology diversified in various directions.
Already existing lines of research discovered the ecological nature of their work.
This is true for vegetation science (Clements 1916; Gleason 1926; Braun-Blanquet
1927; Whittaker 1962), marine ecology (Petersen 1913; Zauke 1989) and limnol-
ogy (Hagen 1992; Golley 1993), all of which had been practised since around 1840
(see for more details Chaps. 19 and 26), and theoretical population ecology (Scudo
1971; Simberloff 1980; Kingsland 1985), which had a longer history. Shortly there-
after, major conceptual progress was marked by the work of C. Elton (1927) and
Tansley (1935).
Phase 6. A second phase of scientific progress occurred after World War II. The
development of “New Ecology” was accompanied by the publication of E.P.
Odum’s textbook in 1953 and the development of the systems concept in ecology
(Worster 1977; Golley 1993). Simultaneously, the work of R. MacArthur supported
the progress of community ecology. Both lines of development are connected by
the contribution of G.E. Hutchinson, who was likewise the supervisor of R.L.
Lindeman (whose work strongly influenced Odum’s work) and R. MacArthur
(Hagen 1992). Both lines culminated in the development of key concepts of modern
ecology (Cherrett 1989). The phase is characterized by growing mathematization
and formalization, turning from descriptive to process and causality orientated
research.
Phase 7. Since the beginning of the 1980s various attempts have been made to unify
ecology based on partly iconoclastic ecology critique (Harper 1982; Peters 1991),
hierarchy theory (Allen et al. 1984) or neutral theories (Hubbell 2001).
Benchmark Papers and Influential Books
Benchmark papers imposed a certain direction on further research (see also Keller
and Golley 2000) since 1920 when, after the foundations of the British and
American ecological societies, indications of professionalization in ecology
became visible (Lowe 1976; Cittadino 1980). Table 7.1 shows an idiosyncratic
overview of benchmark papers.
There are different types of benchmark papers. Gleason (1926) was a concept
paper or rather a self-review. It had no immediate consequences, revival of his ideas
only took place after the publication of Bray and Curtis (1957) and other papers of
the Wisconsin school. Lindeman (1942) published an original research paper
executing conceptual ideas of others (Tansley 1935), subsequently stimulating fur-
ther research. A similar relationship can be found between the work of Volterra
(1926) and Gause’s (1934) book. The subsequent papers of Fisher et al. (1944),
102 G. Wiegleb
Preston (1962) and Connor and McCoy (1979) still form the backbone of community
ecology. Connell and Slatyer (1977) are still unrivalled in the field of succession
theory. Interestingly, since 1980 no important original paper appeared, while hier-
archy theory, scaling theory and theoretical treatment of heterogeneity and spatial
autocorrelation made some progress (see Kolasa and Pickett 1991; Wiegleb 1992;
Palmer and White 1994; Jax et al. 1996).
Many important ideas were first published in books. Table 7.2 shows another
idiosyncratic overview of important ecology books. The first textbook by Warming
(1895: English edition 1909) was replaced by a new textbook only 50 years later
(Odum 1953). A new generation of textbooks (Krebs 1975; Begon et al. 1986)
replaced Odum’s work, which is no longer edited.
Table 7.1 List of benchmark papers
Author Year Keywords
Arrhenius 1921 Species-area relation
Gleason 1926 Individualistic concept
Volterra 1926 Lotka-Volterra equation
Tansley 1935 Ecosystem
Lindeman 1942 Trophic dynamic approach
Egler 1942 Relay floristics
Fisher, Corbet and Williams 1944 Diversity
Novikoff 1945 Levels of organisation
Leslie 1945 Leslie matrix
Watt 1947 Mosaic cycle
Skellam 1951 Random dispersal
Hutchinson 1957 Niche
Bray and Curtis 1957 Ordination
Huffaker 1958 Predator–prey systems
Slobodkin 1960 Energy relations
Connell 1961 Competition
Preston 1962 Species individual relation
MacArthur and Wilson 1963 Island biogeography
Margalef 1963 Ecosystem theory
Yoda et al. 1963 Self-thinning
Paine 1966 Food webs
Bormann and Likens 1967 Biogeochemistry
Porter and Gates 1969 Biophysical ecology
Simberloff and Wilson 1969 Zoogeography
May 1974 Stability
Connell and Slatyer 1977 Succession models
Connor and McCoy 1979 Passive sampling
Harper 1982 After description
Juhász-Nagy and Podani 1983 Spatial processes
Allen, O’Neill and Hoekstra 1984 Hierarchy theory
Kolasa and Pickett 1989 Scaling theory
Legendre 1993 Spatial autocorrelation
1037 A Few Theses Regarding the Inner Structure of Ecology
Elton’s (1927) can likewise be regarded as the necessary counterpart to Warming
in the field of animal ecology. Beyond that it presented conceptual novelties and led
to enhanced empirical research. Small books on seemingly restricted subjects such
as Grime (1979); Box (1981); Tilman (1982) or Hubbell (2001) triggered empirical
research and had impact far beyond their original intensions.
Current Structure of Ecology
Organisms (Systematic and Functional Groups)
Ecology of systematic groups distinguishes between microorganisms (microbial ecology,
geomicrobiology), plants (geobotany, plant ecology; mostly restricted to vascular
plants); and animals (animal ecology, concentrating on vertebrates and insects).
Ecology of man (human ecology) is excluded here as a special case, as it is rather a
hybrid of ecology with social sciences. Classification by Wikipedia (2004) deliberately
includes human ecology in the treatment. All subgroups have their own textbooks and
journals. Departments at universities and funding organisations are orientated towards
the taxonomic approach. Justification of the taxonomic approach is derived from
objective differences between major taxa, e.g. trophic status or mobility. Dodson’s
(1998) “organism approach” also allows functional classifications according to growth
form (tree ecology), mobility type (zooplankton ecology), trophic relations (parasite
ecology) or life span (ecology of long lived organisms). Life form classifications
(e.g. Raunkiaer 1934) played an important role in plant ecology.
Table 7.2 List of important books
Author Year Keywords
Warming 1895 Community
Clements 1916 Succession
Elton 1927 Animal ecology
Gause 1934 Competition
Odum 1953 General textbook
Andrewartha and Birch 1954 Population ecology
MacArthur 1972 Geographic ecology
Harper 1975 Population ecology
Krebs 1975 General textbook
Green 1979 Research method
Grime 1979 Plant strategies
Gates 1980 Biophysical ecology
Box 1981 Growth form and climate
Tilman 1982 Competition
Begon, Harper and Townsend 1986 General textbook
Allen and Hoekstra 1992 Unification
Hubbell 2001 Neutral theory
104 G. Wiegleb
Habitat Types
Ecology of habitat types or locations (Dodson 1998) is divided along the land-water
borderline. Terrestrial ecology includes further subdivisions such as tropical to
polar ecology, desert to forest ecology, or agro-, forest and urban ecology. Soil is
often treated as a separate habitat type (soil ecology). Aquatic ecology deals with
the sea (as does marine ecology) or inland waters (as limnology, hydrobiology, or
freshwater ecology). The latter is, furthermore, strongly divided into lake and river
ecology, sometimes including, sometimes deliberately excluding wetland ecology.
This subdivision can be encountered in denominations of ecological departments,
in the organisation of chapters in ecology textbooks, and in specific journal and
societies devoted to the study of habitat centred ecology. Justification of the habitat
approach is derived from objective differences between habitat types, including
main limiting factors, homogeneity of resource distribution, or driving forces in
community and ecosystem organisation.
Observational Levels Versus Scales
The idea of observational levels is derived from hierarchy theory (Novikoff 1945).
It was introduced by Eugene P. Odum (1959) in the organisation of an ecology
textbook. Observational levels serve for the constitution of ecological units (Jax
et al. 1998). Different levels are treated in own textbooks: Individuals (autecology),
populations (demecology, population ecology), communities (community ecology,
biocoenology, often called “synecology”), ecosystems (ecosystem ecology, system
ecology), and landscapes (landscape ecology). Dodson (1998), following Allen and
Hoekstra (1992) is speaking of perspectives instead of observational levels. Thus
“individual ecology” is separated into physiological ecology and behavioural ecol-
ogy. Physiological ecology may include chemical, molecular or biophysical ecol-
ogy. Exotoxicology, on the other hand, is no subdiscipline of ecology. Even though
dealing with the same substances as physiological or habitat ecology the centre of
interest is the chemical substance and its fate and not any biological entity.
The range of observational levels covered by ecology is shown in Table 7.3.
Levels range from the individual to the biome. Only Allen and Hoekstra (1992)
recognize all levels as ecologically relevant. Some authors (Shelford 1913), in the
tradition of Warming or Klötzli (1989, being strongly influenced by systems think-
ing) regarded one of the levels as truly ecological. One might argue that any state-
ment on the appropriate level of observation in ecology has paradigmatic character.
While some of the observational levels refer to a defined scale (e.g. landscape
ecology is always large-scale, Forman and Godron 1996), others such as population
or community ecology can be carried out on different scales (Allen and Hoekstra
1990). A distinction of ecological approaches according to spatial scales or spatio-
temporal domains was introduced by Delcourt et al. (1983). Different spatial scales
and coinciding temporal scales require different research strategies. In plant ecology
1057 A Few Theses Regarding the Inner Structure of Ecology
this was reflected by the distinction between small-scale “vegetation science” and
large-scale “phytogeography”. Recently, large scale ecology was linked to com-
munity ecology as “macroecology” (Gaston and Blackburn 1999).
Key Concepts
Key concepts had a major influence on the course of research in ecology (see Chap. 2).
Based on an interview of 500 British ecologists on the occasion of the 75th anni-
versary of the British Ecological Society (in 1986), the following key concepts of
ecology, ordered according to organisational levels, were identified. In brackets,
ranking positions are given. Some concepts may appear twice or more (e.g. diversity
includes both as “diversity” and “diversity-stability-relation”).
Table 7.3 Observational levels of ecology
Author Individual Population Community Ecosystem
Landscape,
biome
Begon et al. (1986) X X X – –
Southwood (1977) – X X – –
Shelford (1913) – – X – –
Allen and Hoekstra (1992) X X X X X
Rowe (1961) X – – X –
Odum (1971a) – X X X X
Klötzli (1989) – – – X –
Table 7.4 Research programmes in ecology (revised after Wiegleb 1996)
Level Individuals and
population Community Ecosystem and landscapeYear
1895 – Warming (causal ecology) –
1942 – – Lindeman (trophic-
dynamic approach)
1954 Andrewartha and Birch
(distribution and
abundance)
– –
1967 Harper (Darwinian
ecology)
– Borman and Likens (bio-
geochemical cycles)
1972 – MacArthur (geographical
ecology)
–
1984 Allen, O’Neill and
Hoekstra (hierarchy
theory)
Allen, O’Neill and Hoekstra
(hierarchy theory)
Allen, O’Neill and
Hoekstra (hierarchy
theory)
1988 – Carpenter and Kitchell (trophic
cascades)
Carpenter and Kitchell
(trophic cascades)
1996 – Mooney et al. (biodiversity
and ecosystem function)
Mooney et al.
(biodiversity and
ecosystem function)
106 G. Wiegleb
Population and evolutionary ecology: Life History and Strategies (9; also 12, –
Adaptation), various aspects of Population Dynamics (15, 17, 19), Coevolution
(24, also 34), r and K selection (33).
– Community ecology: Succession (2; also 41, Climax), Competition (5; also 30,
Competitive Exclusion), Niche (6), the Community (8), Species diversity (14;
incl. 35, Diversity-stability Relation), Limiting factors (16), Predation (20, also
21 and 38), Island Biogeography and Species-area Relation (22, 39), Natural
Disturbance (26), Indicator Organisms (29).
– Ecosystem and landscape ecology: The Ecosystem (1), Energy Fluxes (3),
Material Cycling (7), Ecosystem Fragility (10), Food Webs (11, also 31),
Heterogenity (13), Maximum Sustainable Yield (18).
– Nature conservation and environmental protection: Resource protection (4),
Bioaccumulation (23), Habitat Restoration (27), Management of nature reserves (28).
Key concepts are almost equally distributed among the population, community and
ecosystem level. In contrast to the community and the ecosystem, the population is
not regarded as a genuine ecological concept. Ecosystem and community concepts
take the highest ranks (ecosystem, succession, energy flux, competition, niche,
material flux, community and diversity). Some concepts are related to dynamic
processes (succession, material and energy cycling), others to important driving
forces (competition, disturbance). The only applied concept in the Top 10 is
resource protection. Today “resource protection” would surely be replaced by
“sustainability”.
Today, the ecosystem is still important as a general framework for ecological
studies. However, most attempts to discover nontrivial ecosystem properties (emer-
gent properties such as goal functions) failed (see Jørgensen 1992; Müller et al.
1996; Gnauck 2002). Energy and material flux belong to the “trivial” (reductionis-
tic) properties of ecosystems, which need to be studied for the description of the
system but which don’t seem to have any interesting relationship to non-obvious
biotic parameters. Succession is a concept dating back to pre-Linnean times
(Clements 1916). Despite being a prominent concept, succession theory is still
immature. This is due to the fact that long-term observations are lacking for most
community types and many scientific ideas are based on indirect inference (short-
term observations, chrono-sequences, spatial gradients).
Competition is one of the most disputed concepts in ecology. Competition is easy
to produce under experimental circumstances (De Wit 1962), but almost impossible
to observe under natural conditions. Again, indirect inference prevails (e.g. null
models, Harvey et al. 1983; Gotelli and Graves 1996). The importance of the niche
concept is clearly declining and niche theory might be regarded as a part of a com-
petition theory. From my personal point of view, the community itself is and will be
the central concept of ecology, despite the fact that the dispute between the func-
tional community view and the statistical community view is still unsettled.
The concept of diversity has been recently transformed into “biodiversity”.
Most probably it would be voted No. 1 in a current poll. However, inclusion of
nonbiological aspects has created some communication and research problems.
1077 A Few Theses Regarding the Inner Structure of Ecology
The biological part of biodiversity is intimately related to other important
key-concepts such as the species-area relationship. Another concept of growing
importance is disturbance. Disturbance is a more recent concept, even though ideas
of catastrophic events can be traced back to the times of Linnaeus and Warming
(Worster 1977). Main hindrance for the development of disturbance theory is the
fundamental disagreement on the nature of disturbance as defined by Grime (1979)
versus Pickett and White (1985).
Research Programmes
Research programmes have played an important role in ecology (Wiegleb 1996).
Nine research programmes are distinguished in Table 7.3. Partly they were pub-
lished in books, partly in conceptual papers. The first ecological research pro-
gramme advanced by E. Warming referred to the level of the plant community.
Community ecology soon reached zoology (Petersen 1913; Shelford 1913). The
community ecology research programme was later refined and generalized by
MacArthur (1972) treating almost any important community ecology concept. As
the treatment of Hubbell (2001) shows, the process of integrating disparate view on
community ecology and relevant aspects of population and evolutionary ecology is
still unfinished. Hubbelll’s book does not include a truly new research programme.
Lindeman (1942) created a new “trophic-dynamic approach” as a counterpart to
the “static species distribution approach” (comprising classification of communi-
ties, habitat ecology, life and growth form analysis) and the “dynamic species dis-
tribution approach” (including succession research). Both refer to Warming’s
approach and Clements’ dynamic extension. Further ramifications of the trophic
dynamic approach have been described by Golley (1993). The most original one is
that of Bormann and Likens (1967) which might be regarded as a research pro-
gramme in its own right. Some reductionistic approaches (e.g. Odum 1971b) fullfill
the requirement of being a true research programme, but not necessarily one in
ecology. The integration of ecosystem ecology and community ecology, which had
been lost in the times of hard-boiled systems ecology was re-established by
Carpenter and Kitchell (1988), based on an idea of Hairston et al. (1960); Mooney
et al. (1996) made another integrative attempt. Also their idea was not new, but can
be found in many papers since the end of the 1980s. Mooney et al. (1996) reached
popularization, as the time was ripe for a revival of a more elaborate version of the
diversity-stability-hypothesis.
Population ecology for a long time remained divided into studies on animal
(Andrewartha and Birch 1954) and plant populations (Harper 1975). Yet, in marine
ecology a separate line of population and life-history-centred ecology prevailed (Zauke
1989). Despite the obvious effort of Hubbell (2001) the lose ends of “individualistic”
community ecology (Gleason 1926), Darwinian ecology (Harper 1967), individual
based modelling (Wissel 1989), theoretical population ecology (Volterra 1926; Pielou
1969), and life history theory (Stearns 1976) still need further integration.
108 G. Wiegleb
At first glance, hierarchy theory of Allen et al. (1984) does not relate to any of
the preceding traditions. Instead, it tries to unify all possible research programmes
in ecology under one common headline, hierarchy theory (see also Chap. 6). But it
is clearly an offspring of the Wisconsin school of plant ecology, trying to integrate
achievements of modern science and systems theory into community ecology in a
wider sense.
Schools and Traditions
The distinction between research programmes and schools (or “traditions”) assumes
that there are schools within larger research programmes, or even schools outside
of recognizable research programmes. Schools characterise a lower level of scien-
tific integration. Differentiation into schools is well described for systems ecology
(Golley 1993), community ecology (Hubbell 2001) and vegetation science
(Whittaker 1962).
Ecosystems ecology followed quite divers directions, e.g. thermodynamics,
exergy, networks, cybernetics, automaton theory etc. One might distinguish a
Margalef school, an H.T. Odum school, a Jørgensen school etc. (Regier and Rapport
1978; Jørgensen 1992; Golley 1993; Gnauck 2002). In contrast to the assumptions
of Regier and Rapport (1978) none of these schools ever reached the paradigmatic
stage. Ulanowicz (1990) paper can be regarded as the last attempt to save systems
ecology and render Harper’s (1982) and Fenchel’s (1987) criticism obsolete.
Community ecology, differentiated according to the interest in competition as a
major driving force of structuring natural communities (the ghost of competition past,
assembly rules; Diamond 1975; Connor and Simberloff 1979) and to methodological
preferences (e.g. Pielou 1969; Hurlbert 1990; mathematical ecology). Today the situ-
ation of community ecology is more relaxed, even though approaches centred around
null models (Strong 1980; Gotelli and Graves 1996) or neutral theories (Hubbell
2001) tend to undermine the conventional wisdom of community ecology.
Vegetation science has been analyzed by Whittaker (1962). At an early stage,
vegetation science split into conflicting schools based on floristic traditions
(emphasis on species composition: north European, south European, Russian,
British and an American tradition) or physiognomic traditions (emphazising vege-
tation structure or architecture; found in Scandinavia, North America, also in tropi-
cal ecology). Simberloff (1980) assumes that vegetation science differentiated on a
paradigmatic level. However, scientists such as Gleason, Clements and Tansley can
still be seen as members of Warming’s research programme. This is not true for
Central European phytosociology (Braun-Blanquet 1927; see Chap. 21 and also
Chap. 19). The Wisconsin school (Bray and Curtis 1957) founded a new tradition
within an old research programme. Major progress in computer technology and
programming (Gauch 1982, Ter Braak 1988) lead to a relaxion of tensions between
schools and eventually to an intrusion of vegetation scientific methods into the core
of community ecology.
1097 A Few Theses Regarding the Inner Structure of Ecology
Applicability
Distinctions according to applicability can be drawn between theoretical ecology or
basic ecology on the one hand and applied ecology on the other (Dodson 1998). The
terms “theoretical” and “basic” have different connotations. Theoretical ecology is
mostly equated to mathematical ecology (Pielou 1969; Wissel 1989), even though
mathematical models can be applied to practical questions. Basic ecology refers to
any scientific research, the results of which can be transferred to applied sciences
such as conservation biology, agriculture, wildlife management or restoration
ecology (Simberloff 1999). Ecology is a pure or basic natural science. For the appli-
cation of ecological knowledge additional normative elements are needed (see Chap.
26). For the relationship between ecology and its applied sciences such as nature
conservation, the same relationship holds as between physiology and medicine, or
physics and electrical engineering. This distinction is confused in many ecological
writings. As a recent example, Wikipedia (2004) is mentioned. The distinction
between basic and applied is, however, dynamic. Genecology and palaeoecology
have changed recently from theoretical exercises to applied disciplines.
Crossover of ideas lead to the introduction of “biodiversity research” (CBD
1992) including both a basic part (systematics and ecology) and an applied part
(planning and socioeconomy; Wiegleb 2003). This is a reaction to the fact that basic
ecological research is not funded in most countries. In Germany, most funding was
spent on applied ecological research in the context of explaining forest die-back,
mitigating consequences of intensive agriculture, restoration of rivers, lakes, flood
plains, or drainage areas, and the like. Many of these studies yielded interesting
results concerning the case studied. Casually, these studies also yielded concepts of
theoretical interest (e.g. Müller et al. 1996; Hauhs and Lange 1996). An attempt to
proceed in a different way (study applied questions on a solid theoretical basis,
Zauke 1989; Vareschi and Zauke 1993) never received adequate funding and was
closed down before having really started.
Conclusions
Is there really “more to ecological science in the postdescriptive phase than acquiring
the ability to handle unique anecdotal management problems” (Harper 1982)? The
above discussion shows that there are few general laws or rules available in ecology,
that there is no unification in ecology as to aims, methods and conceptual
approaches, and that all attempts to reduce the separate ecologies to one consistent
bulk of knowledge have failed so far. So far, no general agreement as to the question
of Dice (1955) “What is ecology?” has been reached. Progress has been made in
the description of single species or single habitats. However, most of the knowledge
collected cannot be transferred to other species of different life form types or other
habitats under different climatic conditions.
110 G. Wiegleb
We are confronted with the unusual situation that ecology is not characterized
by paradigm formation. “Paradigms” in ecology such as thermodynamics, stoechi-
ometry, or evolution are borrowed from other sciences. We observe a separation of
ecology into simultaneous schools or traditions that behave like paradigms.
Communication among schools does not take place either because it is not intended
(lacking respect for other sub-disciplines) or impossible (language barriers).
I assume that scientific communication has been possible since 1800, even though
the sheer number of new publications nowadays may impose physical boundaries
on information transfer. In the 1930s in the USA, adherents of Clements would not
read any paper by Gleason. Followers of a “physicalistic” school (Odum 1971b;
Jørgensen 1992) would never read papers such as Harper (1967, 1982); Walter
(1973); Stearns (1976) or Krebs and Davies (1993). Followers of the Braun-
Blanquet approach deliberately neglected the whole bulk of Anglo-American com-
munity ecology for decades. Likewise, the important paper of Petersen (1913) was
neglected by terrestrial ecologists. Attempts to bridge the gap between observational
levels (e.g. Gates 1980; Carpenter and Kitchell 1988; Turner 1989; Ulanowicz 1990;
Jones 1995; Bartha et al. 1998) are usually not recognized outside a specialized
group of scientists. Other people might think that Carpenter and Kitchell (1988)
wrote on “lakes”, but their topic is much wider.
Even within its traditions, ecological knowledge is far from being cumulative.
Are we really wiser with respect to the life history of plants than Salisbury (1932)
has been? Good old wisdom is forgotten (e.g. the nonvalidity of crude forms of
the stability-diversity hypothesis, Goodman 1975; Trepl 1995). Money is still
being spent to prove this obsolete hypothesis. Nowadays it is difficult to distin-
guish between a proper research programme and ephemeral fashions. Research in
ecology has become opportunistic. Philosophers, economists, politicians, and
jurists have adopted ecological terms or rather developed their own “creole” or
“pidgin” languages on ecology. The ecologist, e.g. when reading the advertise-
ments of funding organisations, has to understand these texts. The main reason
for this situation is that scientific standards such as recognition of the stochastic
and historic nature of ecological systems (Simberloff 1980; Levins and Lewontin
1980), the necessity of predictive ecology (Harper 1982; Peters 1991), or the
necessity of inter-level reduction of phenomena (Allen et al. 1984; Kolasa and
Pickett 1989; Shrader-Frechette and McCoy 1990) have not yet become common
sense in ecology.
Acknowledgment I thank U. Böring for the critical reading of various versions of the text and
help with the technical preparations of the manuscript.
References
Abbot D (1983) The biographical dictionary of scientists. Biologists. Blond Educational, London
Allen TFH, Hoekstra TW (1990) The confusion between scale-defined levels and conventional
levels of organisation. J Vegetable Sci 1:5–12
1117 A Few Theses Regarding the Inner Structure of Ecology
Allen TFH, Hoekstra TW (1992) Toward a unified ecology. Columbia University Press, Columbia
Allen TFH, O‘Neill RV et al (1984) Interlevel relations in ecological research and management:
some working principles from hierarchy theory. USDA, Forest Service, General Technical
Report RM-110
Andrewartha HG, Birch LC (1954) The distribution and abundance of animals. University of
Chicago Press, Chicago
Anker P (2001) Imperial ecology: the environmental order of the British Empire, 1895-1945.
Harvard University Press, Cambridge
Arrhenius O (1921) Species and area. J Ecol 9:95–99
Bartha S, Czaran T et al (1998) Exploring plant community dynamics in abstract coenostate
spaces. Abstracta Bot 22:49–66
Begon M, Harper JL et al (1986) Ecology: individuals, populations and communities. Blackwell,
Sunderland
Bormann FH, Likens G (1967) Nutrient cycling. Science 155:424–429
Box EO (1981) Macroclimate and plant form: an introduction to predictive modelling in phyto-
geography. Junk, the Hague
Braun-Blanquet J (1927) Pflanzensoziologie. Bornträger, Berlin
Bray JR, Curtis JT (1957) An ordination of the Upland Forest Communities of Southern
Wisconsin. Ecol Monogr 27:325–349
Bremekamp CEB (1952) Linné’s significance for the development of phytogeography. Taxon
2:47–54
Carpenter S, Kitchell JF (1988) Strong manipulations and complex interactions: consumer control
of lake productivity. Bioscience 38:764–769
CBD (1992). from www.biodiv.org.
Cherrett JM (1989) Key concepts: the result of a survey of our members’ opinions. In: Cherrett
JM (ed) Ecological concepts: the contribution of ecology in understanding of the natural
world. Blackwell, Oxford, pp 1–16
Cittadino E (1980) Ecology and the professionalization of botany in America, 1890-1905. Stud
Hist Biol 4:171–198
Clements FE (1916) Plant succession. An analysis of the development of vegetation. Carnegie
Institution of Washington, Washington, DC
Connell JH (1961) The influence of interspecific competition and other factors on the distribution
of the Barnacle Chthamalus stellatus. Ecology 42:710–723
Connell JH, Slatyer RO (1977) Mechnisms of succession in natural communities and their role in
community stability and organization. Am Nat 111:1119–1144
Connor EF, McCoy ED (1979) The statistics and biology of the species-area relationship. Am Nat
113:791–833
Connor EF, Simberloff D (1979) The assembly of species communities: chance or competition?
Ecology 60:1132–1140
De Wit C (1962) On competition. Pudoc, Wageningen
Delcourt HR, Delcourt PA et al (1983) Dynamic plant ecology: the spectrum of vegetational
change in space and time. Q Sci Rev 1:153–175
Diamond JM (1975) Assembly rules of species communities. In: Cody ML, Diamond JM (eds)
Ecology and evolution of communities. Harvard University Press, Cambridge, pp 342–444
Dice LR (1955) What is ecology? Sci Monogr 80:346–351
Dodson S (ed) (1998) Ecology. Oxford University Press, Oxford
DuRietz GE (1957) Linnaeus as a phytogeographer. Vegetatio 6:161–168
Egerton FN (1968) Studies on animal populations from Lamarck to Darwin. J Hist Biol 3:225–259
Egerton FN (1969) Ricardo Bradley’s understanding of biological productivity: a study of eigh-
teenth-century ecological ideas. J Hist Biol 2:391–410
Egerton FN (1973) Changing concepts of the balance of nature. Q Rev Biol 48:322–350
Egerton FN (1976) Ecological studies and observations before 1900. In: Taylor BJ, White TJ (eds)
Issues and ideas in America. University of Oklahoma Press, Oklahoma, pp 311–351
Egerton FN (1977) A bibliographical guide to the history of general ecology and population biology.
Hist Sci 15:189–215
112 G. Wiegleb
Egerton FN (1983) The history of ecology: achievements and opportunities, part one. J Hist Biol
16:259–310
Egler FE (1942) Vegetation as an object of study. Philos Sci 9:245–260
Elton C (1927) Animal ecology. Methuen, London
Fenchel T (1987) Ecology potentials and limitations. International Ecology Institute, Oldendorf
Fisher RA, Corbet AS et al (1944) The relation between the number of species and the number of
individuals in a random sample of an animal population. J Anim Ecol 12:42–58
Forman RTT, Godron M (1996) Landscape ecology. Wiley, New York
Gaston KJ, Blackburn TM (1999) A critique for macroecology. Oikos 84:353–368
Gates D (1980) Biophysical ecology. Springer, New York
Gauch HG (1982) Multivariate analysis in community ecology. Cambridge University Press,
Cambridge
Gause GF (1934) The struggle for existence. Williams & Wilkins, Baltimore
Gleason HA (1926) The individualistic concept of the plant associations. Bull Torrey Bot Club
53:7–26
Gnauck A (2002) Automatentheorie in der Ökologie. In: Gnauck A (ed) Systemtheorie und
Modellierung von Ökosystemen. Physica, Berlin, pp 32–48
Golley FB (1993) A history of the ecosystem concept in ecology: more than the sum of the parts.
Yale University Press, New Haven
Goodland RJ (1975) The tropical origin of ecology: Eugen Warming´s Jubilee. Oikos
26:240–245
Goodman D (1975) The theory of diversity-stability relationships in ecology. Q Rev Biol
50:237–266
Gotelli NJ, Graves GR (1996) Null models in ecology. Smithsonian Institution Press, Washington,
DC
Green RH (1979) Sampling design and statistical methods for environmental biologists. Wiley-
Interscience, New York
Grime JP (1979) Plant strategies and vegetation processes. Wiley, Chichester
Hagen JB (1992) An entangled bank: the origins of ecosystem ecology. Rutgers University Press,
New Brunswick
Hairston N, Frederick G et al (1960) Community structure, population control, and competition.
Am Nat 94:421–425
Harper JL (1967) A Darwinian approach to plant ecology. J Ecol 55:247–270
Harper JL (1975) Population ecology of plants. Academic, New York
Harper JL (1982) After Description. In: Newmann EI (ed) The plant community as a working
mechanism. Blackwell, London, pp 11–25
Harvey PH, Colwell RK et al (1983) Null models in ecology. Annu Rev Ecol Syst 14:189–211
Hauhs M, Lange H (1996) Ecosystem dynamics viewed from an endoperspective. Sci Total
Environ 183:125–136
Hubbell SP (2001) The Unified Neutral Theory of biodiversity and biogeography. Princeton
University Press, Princeton
Huffaker CB (1958) Experimentals studies on predation: dispersion factors and Predator-Prey
Oscillations. Hilgardia 27:343–383
Hurlbert SH (1990) Spatial distribution of the montane unicorn. Oikos 58:257–271
Hutchinson GE (1957) Concluding remarks, population studies: animal ecology and demography.
Cold Spring Harb Symp Quant Biol 22:415–427
Jahn I, Löther R et al (eds) (1985) Geschichte der Biologie. Theorien, Methoden, Institutionen und
Kurzbiographien. Fischer, Jena
Jansen AJ (1972) An analysis of “Balance of Nature” as an ecological concept. Acta Biotheor
21:86–114
Jax K, Jones CG et al (1998) The self-identity of ecological units. Oikos 82:253–264
Jax K, Potthast T et al (1996) Skalierung und Prognoseunsicherheit bei ökologischen Systemen.
Verhandlungen der Gesellschaft für Ökologie 26:527–535
Jones CG (ed) (1995) Linking species and ecosystems. Chapman & Hall, New York
1137 A Few Theses Regarding the Inner Structure of Ecology
Jørgensen SE (1992) Integration of ecosystems theories: a pattern. Kluwer, Dordrecht
Juhász-Nagy P, Podani J (1983) Information theory methods for the study of spatial processes and
succession. Vegetatio 51:129–140
Keller DR, Golley FB (eds) (2000) From science to synthesis. Readings in the foundational con-
cepts of the science of ecology. University of Georgia Press, Athens
Kingsland SE (1985) Modelling nature. Episodes in the history of population ecology. University
of Chicago Press, Chicago
Klötzli F (1989) Ökosysteme. Fischer, Stuttgart
Kolasa J, Pickett STA (1989) Ecological systems and the concept of biological organisation. Proc
Natl Acad Sci USA 86:8837–8841
Kolasa J, Pickett STA (eds) (1991) Ecological heterogeneity. Ecological studies. Springer, New York
Krebs CJ (1975) Ecology. The experimental analysis of distribution and abundance. Harper
Collins, New York
Krebs CJ, Davies NB (1993) An introduction to behavioural ecology. Blackwell, London
Krebs CJ (1995) Ecology. The experimental analysis of distribution and abundance. Harper
Collins, New York
Kuhn TS (1976) Die Struktur wissenschaftlicher Revolutionen. Suhrkamp, Frankfurt
Küppers G, Lundgreen P et al (1978) Umweltforschung - die gesteuerte Wissenschaft? Eine
empirische Studie zum Verhältnis von Wissenschaftsentwicklung und Wissenschaftspolitik.
Suhrkamp, Frankfurt
Lakatos I (1978) Die Geschichte der Wissenschaft und ihre rationale Rekonstruktion. In:
Diederich W (ed) Theorien der Wissenschaftsgeschichte. Suhrkamp, Frankfurt, pp 55–119
Legendre P (1993) Spatial autocorrelation: trouble or new paradigm? Ecology 74:1659–1673
Leslie PH (1945) On the use matrices in certain population mathematics. Biometrika
33:183–212
Levins R, Lewontin R (1980) Dialectics and reductionism in ecology. Synthese 43:47–78
Lindeman RL (1942) The trophic-dynamic aspect of ecology. Ecology 23:339–418
Lowe PD (1976) Amateurs and professionals: the institutional emergence of British plant ecology.
J Soc Bibliogr Nat Hist 7:517–535
Lussenhop J (1974) Victor Hensen and the development of sampling methods in ecology. J Hist
Biol 7:319–337
MacArthur RH (1972) Geographical ecology. Harper & Row, New York
MacArthur RH, Wilson EO (1963) An equilibrium theory of insular zoogeography. Evolution
17:373–387
Mägdefrau K (1992) Geschichte der Botanik. Leben und Leistung großer Forscher. Fischer,
Stuttgart
Margalef R (1963) On certain unifying principles in ecology. Am Nat 97:357–374
May RM (1974) Biological populations with non-overlapping generation: stable cycles, and
chaos. Science 186:645–647
May RM, Seger J (1986) Ideas in ecology. Am Sci 74:256–267
Mayr E (1984) Die Entwicklung der biologischen Gedankenwelt. Vielfalt, Evolution und
Vererbung. Springer, Berlin
McIntosh RP (1985) The background of ecology: concept and theory. Cambridge University
Press, Cambridge
McIntosh RP (1987) Pluralism in ecology. Annu Rev Ecol Syst 18:321–341
Mooney HA, Cushman JH et al (eds) (1996) Functional roles of biodiversity – A global perspec-
tive. Wiley, Chichester
Müller F, Fränzle O et al (1996) Modellbildung in der Ökosystemanalyse als Integrationsmittel
von Empirie, Theorie und Anwendung – eine Einführung. EcoSys 4:1–16
Nelson G (1978) From candolle to croizat: comments on the history of biogeography. J Hist Biol
11:269–305
Novikoff AB (1945) The concept of integrative levels and biology. Science 101:209–215
Odum EP (1953) Fundamentals of ecology. Saunders, Philadelphia
Odum EP (1959) Fundamentals of ecology. Saunders, Philadelphia
114 G. Wiegleb
Odum EP (1971a) Fundamentals of ecology. Saunders, Philadelphia
Odum EP, Odum HT (1953) Fundamentals of ecology. Saunders, Philadelphia
Odum HT (1971b) Environment, power and society. John Wiley & Sons, London
Paine RT (1966) Food web complexity and species diversity. Am Nat 100:65–75
Palmer MW, White PS (1994) Scale dependence and the species-area relationships. Am Nat
144:717–740
Peters RH (1991) A critique for ecology. Cambridge University Press, Cambridge
Petersen CGJ (1913) Valuation of the sea II. The animal communities of the sea bottom and their
importance for marine zoogeography. Rep Danish Biol Stat 21:1–44
Pickett STA, White PS (1985) Patch dynamics – a synthesis. In: Pickett STA, White PS (eds)
The ecology of natural disturbance and patch dynamics. Academic, San Diego, pp 371–384
Pielou EC (1969) An introduction to mathematical ecology. Wiley Interscience, New York
Porter WP, Gates DM (1969) Thermodynamic equilibria of animals with environment. Ecol
Monogr 39:224–244
Preston FW (1962) The canonical distribution of commonness and rarity. Ecology
43(185–215):431–432
Querner H (1980) Das teleologische Weltbild Linne’s - Observationes, Oeconomia, Politia. Veröff
Joachim Jungius-Ges Wiss Hamburg 43:25–49
Raunkiaer C (1934) The life forms of plants and statistical plant geography. Clarendon, Oxford
Regier HA, Rapport DJ (1978) Ecological paradigms, once again. Bull Ecol Soc Am 59:2–6
Reise K (1980) Hundert Jahre Biozönose. Die Evolution eines ökologischen Begriffes.
Naturwissenschaftliche Rundschau 33:328–335
Rowe JS (1961) The level-of-integration concept and ecology. Ecology 42:420–427
Salisbury EJ (1932) The East Anglian flora. Trans Norfolk Norwich Natl Soc 13:191–263
Scudo F (1971) Vito Volterra and theoretical ecology. Theor Popul Biol 2:1–23
Shelford VE (1913) Animal communities in temperate America as Illustrated in the Chicago
region. Bulletin of the Geographical Society of Chicago, Chicago
Shrader-Frechette K, McCoy E (1990) Theory reduction and explanation in ecology. Oikos
58:109–114
Simberloff D (1980) A succession of paradigms in ecology: from essentialism to materialism and
probabilism. Synthese 43:3–39
Simberloff D (1999) The role of science in the preservation of forest biodiversity. Forest Ecol
Manage 115:101–111
Simberloff D, Wilson EO (1969) Experimental zoogeography of islands: the colonization of
empty islands. Ecology 50:278–296
Skellam JG (1951) Random dispersal in theoretical populations. Biometrika 38:96–218
Slobodkin LB (1960) Ecological energy relationships at the population level. Am Nat
44:213–236
Southwood TRE (1977) Ecological methods. Chapman & Hall, London
Stauffer RC (1960) Ecology in the long manuscript version of Darwin’s origin of species and
Linnaeus’ Oeconomy of nature. Proc Am Philos Soc 104:235–241
Stearns SC (1976) Life history tactics: a review of ideas. Q Rev Biol 51:3–47
Strong D (1980) Null hypothesis in ecology. Synthese 43:271–285
Tansley AG (1935) The use and abuse of vegetational concepts and terms. Ecology 16:284–307
Ter Braak CJF (1988) CANOCO – A FORTRAN program for canonical community ordination
by [Partial] [Detrended] [Canonical] correspondence analysis and redundancy analysis
(Version 2.1). GLW, Wageningen
Tilman D (1982) Resource competition and community structure. Princeton University Press,
Princeton
Toulmin S (1978) Kritik der kollektiven Vernunft. Suhrkamp, Frankfurt
Trepl L (1987) Geschichte der Ökologie. Athenäum, Frankfurt
Trepl L (1995) Die Diversitäts-Stabilitäts-Diskussion in der Ökologie. Berichte ANL, Beiheft
12:35–49
1157 A Few Theses Regarding the Inner Structure of Ecology
Turner MG (1989) Landscape ecology: the effect of pattern on process. Annu Rev Ecol Syst
20:171–197
Ulanowicz RE (1990) Aristotelian causalities in ecosystem development. Oikos 57:42–48
Vareschi E, Zauke GP (1993) Entwicklung eines theoretischen Konzepts zur Ökosystemforschung
im Wattenmeer. UBA-Texte 47/39:1–142
Volterra V (1926) Fluctuations in the abundance of a species considered mathematically. Nature
118:558–560
Vorzimmer P (1965) Darwin´s ecology and its influence upon his theory. Isis 56:148–155
Walter H (1973) Allgemeine geobotanik. Ulmer, Stuttgart
Warming E (1895) Plantesamfund, grundträk af den Ökologiske plantegeografi. Philipsens Forlag,
Copenhagen
Watt AS (1947) Pattern and process in the plant community. J Ecol 35:1–22
Whittaker RH (1962) Classification of natural communities. Bot Rev 28:1–239
Wiegleb G (1989) Explanation and prediction in vegetation science. Vegetatio 83:17–34
Wiegleb G (1992) Explorative Datenanalyse und räumliche Skalierung – eine kritische Evaluation.
Verhandlungen der Gesellschaft für Ökologie 21:327–338
Wiegleb G (1996) Konzepte der Hierarchietheorie in der Ökologie. In: Mathes K, Breckling B,
Ekschmitt K (eds) Systemtheorie in der Ökologie. Ecomed, Marburg, pp 7–24
Wiegleb G (2003) Was sollten wir über Biodiversität wissen? Aspekte einer angewandten
Biodiversitätsforschung. In: Weimann J, Hoffmann A, Hoffmann S (eds) Messung und
Bewertung von Biodiversität: mission impossible? Metropolis, Marburg, pp 151–178
Wilson DS (1988) Holism and reductionism in evolutionary ecology. Oikos 53:269–273
Wissel C (1989) Theoretische Ökologie. Springer, Berlin
Worster D (1977) Nature´s economy. The roots of ecology. Sierra Club Books, San Francisco
Wuketits FM (1995) Evolutiontheorie. Historische Vorraussetzungen, Positionen, Kritik. Wiss.
Buchgesellschaft, Darmstadt
Yoda K, Kira T et al (1963) Self-thinning in overcrowded pure stands under cultivated and natural
conditions. J Biol Osaka City Univ 14:107–129
Zauke GP (1989) Konzeptionelle Überlegungen für einen Forschungsschwerpunkt “Wattenmeer”
aus der Sicht der theoretischen Ökologie. UBA-Texte: 253–264
117
The Field of Knowledge “Ecology”
In many societies a growing consensus has arisen about the importance of ecological
knowledge: It helps us to address some of the most pressing problems we face at both
global and regional level by providing research and effective management options to
deal with global warming, diminishing natural resources and the deterioration of
soils and water resources. Debates about natural disasters, about the purity of natural
things, and about the perceived crisis of the nature-culture relationship in general all
revolve around the role and the importance of ecological knowledge in social pro-
cesses and in negotiations about the kind of nature with and in which we wish to live.
For this reason it seems not only worthwhile but also essential, in a sense, to take a
closer look at the logical and disciplinary construction of ecological knowledge from
a philosophy of science perspective. Inversely, for the philosophy of science ecology
is an interesting field having identified as one future perspective that it is important
to link general philosophy of science with special philosophies of science in a more
fruitful way. This is the framing I have in mind when I propose in the following to
explore a distinct epistemological approach for ecological knowledge.
First of all, when “philosophy of science deals with the foundations and the meth-
ods of science” its attention is most often focused on the foundations and methods
of a discipline.1 This already entails the expectation that scientific knowledge needs
to be developed in a distinct institutional setting, in one that is in some sense a
“closed society”, with its own language and customs. This is certainly true of scien-
tific disciplines such as physics or astronomy, which have an age-long tradition and
a similarly long tradition of philosophical reflection. But there are other epistemic
domains whose foundations and methods cannot be described adequately by fram-
ing them as a discipline in this sense. They should rather be conceived as a field of
Chapter 8
Dynamics in the Formation
of Ecological Knowledge
Astrid Schwarz
1 This is a widespread characterization of philosophy of science – for instance in the call for paper
for a conference in Tilburg in 2010 organized by Stefan Hartmann and Paul Griffiths on The
Future of Philosophy of Science.
A. Schwarz (*)
Institute of Philosophy, Technische Universität Darmstadt, Schloss, 64283, Darmstadt, Germany
e-mail: schwarz@phil.tu-darmstadt.de
A. Schwarz and K. Jax (eds.), Ecology Revisited: Reflecting on Concepts, Advancing Science,
DOI 10.1007/978-90-481-9744-6_8, © Springer Science+Business Media B.V. 2011
118 A. Schwarz
knowledge that is scattered between different academic disciplines and ultimately
even beyond the academic context itself. I propose, first, that ecology be understood
in the sense of a field of knowledge rather than as a discipline; and second, I propose
an epistemological description to encompass this patchwork field. The theories in
this field move back and forth between three basic conceptions and it is this produc-
tive movement, I argue, that stabilizes ecological knowledge.
At the end of this chapter will be offered another more general perspective that
is a discussion on the relationship between the philosophy of nature and the phi-
losophy of science in view of the field of ecological knowledge. I will provide some
reflections on how the two might be linked and how this could help to better under-
stand the plurality in the field.
The starting point for all these considerations was the finding that one and the
same ecological research object is “seen” in very different ways and that it may be
described simultaneously in different theories and narratives: Organisms in a lake,
for example, can be communities, societies or merely assemblages, depending on
how strongly their mutual interconnection is seen to be and how necessary their
incorporation into their environment. These organisms may have predominantly
friendly or indifferent “neighbourly” relations, or they are hostile towards one
another; the resources available to the organisms exist in unlimited quantity, or they
are described as being permanently scarce; and, finally, organisms themselves can
be conceived of principally as a unit of production, as a storage container or as a
unit of selection.
A good example of an assemblage in terms of a community might be the recent
www-model of a forest (Wiemken and Boller 2002), which describes the rhizomic
system of plants and fungi in a forest. The wood wide web connects all individuals
to one another in a mainly cooperative way and thus enables an exchange of energy,
nutrients and even genetic information in the community. To illustrate an assem-
blage in terms of a society, I have chosen one of the very first descriptions of a
lacustrian system, the microcosm lake, given in 1897 by Stephen Forbes, an
American zoologist who originally started out working in the field of entomology.
Forbes wonders if the “system of life is such that a harmonious balance of conflict-
ing interests has been reached where every element is either hostile or indifferent
to every other, may we not trust much to the outcome where, as in human affairs,
the spontaneous adjustments of nature are aided by intelligent effort, by sympathy,
and by self-sacrifice?” He then describes certain pelagic forms in a lake “often
exquisitely transparent, and hence almost invisible in their native element” which
seems to “protect them (perfectly) against their enemies”. But, “with an ingenuity
in which one may almost detect the flavor of sarcastic humor, Nature has turned
upon these favored children and endowed their most deadly enemies with a like
transparency, so that wherever the towing net brings to light a host of these crystal-
line Cladocera, there it discovers also swimming, invisible, among them, a lovely
pair of robbers and beasts of prey” (1897, p. 545).
Clearly, we are confronted here with different stages of precision of scientific
concepts. Following Rudolf Carnap, one might say that Forbes’ system of life is
still in the classificatory stage, whereas the wood wide web model has already
1198 Dynamics in the Formation of Ecological Knowledge
encompassed the comparative stage and is now in a qualitative stage. This is not
terribly surprising, as there are roughly 100 years of conceptual evolution between
the two. But what might seem surprising is that the conceptual scheme of describing
an assemblage of individuals of plants or animals appears to be quite robust: from
the very beginning of ecological thinking, there have been communities with more
competitive and hostile relations among one another, held together by a contract;
and at the same time, there have been communities where more friendly relations
dominate and/or the reference to a superordinate unity is of utmost importance
(Clements’ superorganism). The idea of the basic conceptions is to capture these
different orientations by describing a connection between the philosophy of nature
involved in a basic conception and the formation of scientific concepts, thus in a
philosophy of science that is interested in the semantic, pragmatic and cognitive
evolution of concepts.
Why Basic Conceptions?
To presuppose this kind of “orientation” is not new. Certainly one could say that
Imre Lakatos’s “hard core” contains also implicit ideas about nature, and this is
certainly the case with the “Naturcharakter” the nature character -, a rather phe-
nomenological concept proposed by Gernot Böhme, which is at work behind the
scenes of the epistemic operations which ascribe meaning to nature. Thus, this
paper argues that ecological plurality is shaped in a distinct way and can be con-
ceptualized as a structure consisting of three so-called basic conceptions. Each of
the three characterizes a particular historical field of knowledge that embraces
practices and theories about living beings in their environments. Over time, basic
conceptions are flexible, they show a dynamic behaviour that is described as an
oscillation. This triadic conceptual system is a suggestion for a dynamic conceptu-
alization of ecological knowledge.
A basic conception is basical in the sense that most ecological theories can be
integrated in one of the three basic conceptions. The basic conceptions embody an
implicit idea about nature that is embedded in the structure and formation of theo-
ries and thus also in the structure and formation of knowledge. The basic concep-
tion operates as a kind of vantage point that organises concepts and theories.
Why Three of Them?
The seductive power of triades is well-known especially when it comes to Hegelian-
Marxist dialectic. It promises a kind of logical and also historical stability that has
a dangerous flipside. Obviously the danger of a dialectical approach lies in the
authoritarian attitude and also in the inexorability of the process. However, it seems
possible to escape these dangers as Lakatos has demonstrated, who was claimed by
120 A. Schwarz
his friend Paul Feyerabend to be a “big bastard – a Pop-Hegelian Philosopher born
from a Popperian father and an Hegelian mother.” (Motterlini 2001, p. 1).
The triade presented here is rather based historically, but nevertheless claims to
give an analytical tool at hand to describe the plurality of ecological theories and
concepts in a heuristically productive and adequate way. The basic conceptions do
not follow one another in the sense of a dialectical scheme, there is no synthetic
position and the dynamic of the basic conception is rather driven by competition to
advance theories or research programmes. These multiple research programmes are
not necessarily related to each other; thus concepts and theories used in the field
might be incommensurable.
Basic Conceptions and Pluralism
The observation that ecology is a science characterized by plurality has become
almost commonplace (Cooper 1996; Kiester 1980; McIntosh 1987; Shrader-
Frechette and McCoy 1994; Haila and Taylor 2001). The reasons given for this plu-
ralism are manifold: Ecology encompasses a variety of epistemological positions
and grapples with a multiplicity of ontological relations. It has often been assumed
by philosophers – and indeed by ecologists themselves – that ecology is an imma-
ture and impure science that ought better to be broken down into separate fields of
knowledge. Even worse, some assume that ecology is just bad science and the idea
of describing ecology as a plural science has frequently been judged as misguided.
This is the case in philosophy2 as well as in ecology itself (see, for example,
Roughgarden 1984; Peters 1991). If we wish to reject such assumptions, it follows
that we need to recognize the partiality of knowledge as important and useful and,
subsequently, to feed it into a different framework that appreciates scientific
plurality.
From the point of view of philosophy, this prompts the question of how such a
framework – one that is open to pluralism and is not constrained by a strong com-
mitment to unified science or even monism – can be adequately conceptualized.
“The multiplicity of approaches is usefully addressed not by comparative evaluations
directed at selecting the uniquely correct one, but by appreciating the partiality of
each. […] In concert, they [the approaches] constitute a nonunifiable plurality of
partial knowledges” (Longino 2006, p. 127). According to this conceptualization,
it is not only admissable to produce partial knowledge, it is also acknowledged
and accepted that these partial knowledges do not operate in parallel, as in a neat
division of labour where every question is approached with its own distinctive set
of methods.
2 See for instance Haila and Taylor (2001, p. 93): “Ecology has received relatively slight attention
among non-biologist scholars, including philosophers, who interpret and comment the life
sciences.”
1218 Dynamics in the Formation of Ecological Knowledge
The present study aligns itself with Longino’s comments and with arguments for
greater tolerance of different knowledge claims and interrelations within disciplines
(conceptual, causal, model-driven, data-driven etc.). It also represents an appeal for
acceptance of different linguistic forms in the sciences and for the permissibility of
different philosophical accounts of scientific methodology. It thus adopts the prag-
matic approach proposed by Rudolf Carnap to learn
from the lessons of history. Let us grant to those who work in any special field of investiga-
tion the freedom to use any form of expression which seems useful to them; the work in
the field will sooner or later lead to the elimination of those forms which have no useful
function. Let us be cautious in making assertions and critical in examining them, but toler-
ant in permitting linguistic forms. (Carnap 1956, p. 40)
Plurality in Ecology
The present approach argues that the existence of a plurality of theories (or pro-
grammes) can have a positive impact because it allows for greater logical flexibility
and thus for more explanatory power as well. On this point, Nancy Cartwright
(1999) has shown that, if anything, the search for explanatory unity detracts from
the search for truth. In a similar vein, Shrader-Frechette and McCoy (1994) argue
in the course of their attempt to develop a philosophical vocabulary adequate to
partial knowledges that, in ecology, the main method used for linking data with a
hypothesis is not the classical deductive scheme but rather (in case studies, for
instance) a variety of logic they call “informal inferences”. Cooper (1998) suggests
a three-fold scheme in which theoretical principles, phenomenological patterns and
causal generalizations are the basic forms of generalization in ecology. This consti-
tutes a philosophical taxonomy which, as Cooper points out, should not be taken as
a rigorous or categorical classification; instead, it should function as an aid to dis-
tinguishing the different modes of investigation (model-driven or data-driven ecology,
for example) and acknowledging their varied generalizations while not dismissing
the possibility that laws may exist in ecology.
The various regions in the taxonomic space […] are all more or less occupied. There is a
great deal of variation in scope and reliability among ecological generalizations. […] (I)f
laws are what philosophy of science has tended to take them to be, then there are no laws
in biology. But that does not mean that everything in biology is equally contingent. (Cooper
1998, p. 582, 584)
Thus all these authors share an unease toward any logical or methodological unity
in science. Instead, they support the idea of different epistemological strategies that
can be described in a philosophically sound way. In ecology these epistemological
strategies range from experimental studies in the lab, through real-world experi-
ments, quasi-experimental studies and case studies, to purely observational studies
in the field.
Taking for granted a plurality of methods and knowledge in ecology, this study
argues that ecological plurality is shaped in a distinct way and can be conceptualized
122 A. Schwarz
as a structure consisting of three basic conceptions, namely “energy”, “niche” and
“microcosm”. These three are conceived of as existing in a rather antagonistic rela-
tion to one another, each of them typifying a particular historical field of historical
knowledge which embraces experimental practices and theories about living beings
in their environments. Over time, all three basic conceptions are flexible; they dis-
play a dynamic behaviour that is described in the following as “oscillation”. The
triadic structure is additionally seen as having systematic implications, in that it
functions as a kind of pattern. This pattern is informed by a distinctive conceptual-
ization of modernity as described from a philosophy of history perspective.
Depending on what political or societal exigencies are at stake (though also as a
result of its internal dynamic), a research programme from the sphere of the
“microcosm” conception may move into the domain of “energy”. It is assumed,
then, that there are relations of competition and dominance between the basic con-
ceptions, such that any one basic conception might push another into the back-
ground but is never able to eliminate it entirely. Instead, a permanent oscillation
between the conceptions enables stability to be attained, without the conceptions
ever becoming unified.3
This may sound rather schematic for now, but this monochrome blueprint will
acquire more colour as examples are provided from the context of theory building
in early ecology. This will be preceded, though, by a section in which a few aspects
of the philosophy of science debate about unity versus plurality in the sciences are
presented. The expectation is that the conceptualization of the basic conceptions
may benefit from this debate – and perhaps, conversely, the pluralism debate may
acquire an additional facet by way of a relatively obscure field of study in the phi-
losophy of science.
Excursus on Plurality and Lawlikeness in Ecology and Biology
Viewing the unity of “science” (in the singular) as a measure of its goodness has
long since ceased to be a commonplace topos in the philosophy of science. The
capacity to think of the sciences in terms of a plurality in both theory and practice
is not only acknowledged as useful, it is now encouraged and indeed required in
this field. An increasing number of proposals – themselves pluralistic – relating to
scientific pluralism have been put forward over the last 20 years. Among their more
well-known exponents are Nancy Cartwright, Ian Hacking, Helen Longino, Alan
Richardson, while their forerunners are often said to include Patrick Suppes, Alfred
North Whitehead, and William Whewell; even Karl Popper and Paul Feyerabend
agreed about this point that “‘theoretical pluralism’ is better than ‘theoretical
monism’” (Lakatos 1972, p. 135).
3 This is explicated in detail later in this essay in the language of the Lakatosian research
programmes.
1238 Dynamics in the Formation of Ecological Knowledge
In the philosophy of the biological sciences (including ecology) special attention
is given to plurality in the sense of the difference and diversity of research objects
and interrelationships. It is this that makes biological research so attractive. This
applies above all to those biological disciplines which are concerned less with
nomothetic than with idiographically constituted knowledge, that is, those areas of
knowledge that are not so much about theory or physical laws in general (as in
molecular biology or evolutionary biology) as about providing an appropriate (in
philosophy of science terms) description of case studies. The issue is thus one of
understanding real world experiments and quasi-experiments, of model-based versus
data-based methods and modes of representation.
The struggle to formulate an appropriate description of this difference and diver-
sity of objects and interrelations has been a part of biological sciences since they
emerged as a field of academic knowledge. What distinguishes the law of gravity
from talk of a law governing the cycle of organic substance, from the basic law of
biocoenosis or from Mendel’s laws? This question of what distinguishes ecological
or biological regularities from physical or chemical regularities is one of determin-
ing and evaluating exceptions and limits, the single and the individual. Also at stake
in this question, though, is a natural philosophy, one which carries with it a claim
to universality and laws in nature. As both ecology and biology grew stronger in
science and society – particularly during the second half of the twentieth century –
and as philosophy of science began to be applied to biological disciplines, contro-
versies arose as to the existence and status of biological (or ecological) laws.
The possibility of biological laws is ruled out from the start when the concept of
(a physical, natural) law is conceived in terms of traditional philosophy of science
and is identified with physics as the ideal science. In this scheme, a natural law
must satisfy three conditions: it must apply at all times and in every place; it must
not contain any reference to individual names; and finally, there must be no excep-
tions. Smart (1963, p. 52) names a fourth condition which amounts to a fourth
argument against the law-based character of generalizations in biology, namely the
role of coincidence. This, so he asserts, can never be completely eliminated in biol-
ogy, which is why “biology is physics and chemistry plus natural history”.4
John Beatty (1995) pursues a similar line of argument to the extent that he, too,
believes there can be no laws in biology on account of the evolutionary contingency
thesis. Wherever it is possible to identify laws in biology, these are not “biological”
laws but ultimately physical or chemical laws. The evolutionary process generates
a situation of “high-level contingency” which ultimately prevents a species from
behaving exactly the same way if “the experiment” were to be repeated – even if
the same environmental conditions prevailed.
This has been countered by the argument that there certainly can be laws in
biology – or at least law-like structures. Martin Carrier, for instance, argues that the
extent to which laws underlie facts is more a matter of degree than of principle:
the concept of fitness, for example, enables us to explain certain aspects of the
4 Smart also stresses that generalizations in biology should be understood in a similar way to those in
the field of technology: both are at the mercy of historical contingency (Schweitzer 2000, p. 369).
124 A. Schwarz
evolutionary process, and the so-called Lotka-Volterra model in ecology makes it
possible to predict the development of a population – within a well-defined range.
The crucial point is to tolerate the idea that these laws
apply to a variety of physically distinct cases. They express features that are quite differ-
ently realized physically. For this reason supervenient concepts are apt to capture general
traits inaccessible on the purely physical level […]. I maintain, biology and the physical
sciences are in the same ballpark. They both contain laws. (Carrier 1995, pp. 92, 97)
What is now virtually uncontested is that this debate about the concept of laws is inex-
tricably linked with a struggle over hierarchies and the power of knowledge within the
sciences. Similarly, most would agree that the debate about realism went hand-in-hand
with a tendency to confuse the level of description with that being described and thus
to “naturalize” the concept of laws. Philosophically, however, this is not a legitimate
move. On this issue, philosopher of science Michael Hampe notes:
The search for syntactic, semantic or ‘architectural’ criteria for law-like regularity is
always a search within the domain of instruments of description, not in that of the aspect
of nature being described. […] One might well say that no philosophically serious attempts
are nowadays made anymore to establish the natural regularity of nature on the basis of an
understanding of nature that is somehow ‘above’ the sciences – which, after all, have
always postulated certain laws of nature.5
This remark concerning the naivety and ignorance of essentialist reasoning is
echoed in a rather laconic remark made by theorist of biology Tim Allen and col-
leagues, who call on ecologists to adapt their narratives to the postmodern world in
which we already live: “The material world will not tell you what decisions you
must make as a scientist … Instead of retreating to naïve objectivism, scientists
need to adapt to a postmodern age by becoming conscious of the significance of
their narratives” (Allen et al. 2001, p. 484).6
To seek to establish laws of nature on the basis of nature itself is philosophically
impossible; as a result, laws lose their normative power as unifying and indeed
necessary instruments of description to certify the scientific character of a discipline.
This opens up a space for other concepts and “narratives” and, not least, for a different
philosophy of nature.
In the 1970s, philosopher of biology Michael Ruse argues against identifying the
concept of laws with physics, as this, he says, relegates biology to the second
5 Hampe 2000, p. 250 f. He also draws attention to the fact that in the “strategy of legitimizing laws
through more general laws, it is not possible to eliminate contingency. Any legitimation of natural
laws has to appeal to something other than laws.” The question of why laws of nature apply, though,
essentially depends, as Hampe points out, on the question of legitimation. God as lawmaker is an
inadmissable option to date; and analytical philosophy of science has no general criterion to offer
either for laws that would enable a distinction to be made between “legitimate” laws and universal
statements which “make an illegitimate claim to the mantle of laws” (see also Giere (1995) for an
extended discussion from a skeptical perspective on the laws of nature). If not otherwise marked,
translation of citations was done by Kathleen Cross.
6 Allen et al. are not only taking ecologists to task here on account of their alleged naive realism;
they are also drawing attention to the problem of underdetermination.
1258 Dynamics in the Formation of Ecological Knowledge
league of science from the start. He also argues, though, that physical laws,
themselves contain exceptions and proper nouns and that Mendel’s Laws most
certainly can be subjected to a process of independent examination (Ruse 1973).
Kenneth Waters (1998) is also wary of the concept of law. He explores a conceptu-
alization of biological generalizations in which a distinction is drawn between two
types of generalization: The first refers to the distribution of traits among biological
entities such as populations or groups, while the second describes dispositions of
causal regularities.7 The trade-off in this distinction is that the evolutionary contin-
gency hypothesis only applies to some biological generalizations, mainly the first
type, that of distribution. However, generalizations about biological regularities can
be made for distinct system classes which are independent in time and space.8
The fact that contingency does not always play a role in ecology is a point also
emphazised by Gregory Cooper. His concern is to find a mechanism for recognizing
degrees of contingency in ecological generalizations. In the end, he suggests that
“the attempt to partition generalizations into the two categories of laws and non-
laws should be abandoned in favor of the concept of nomic force, […] which
recognizes that nomicity in biology comes in degrees and over restricted domains”
(Cooper 1996, p. 33 f.). Similarly, Sandra Mitchell argues that the dichotomous
oppositions “law vs accident” and “necessity vs contingency” are, if anything, an
obstacle that impoverishes the conceptual framework. Such an approach, she says,
“obscures much interesting variation in both the types of causal structures studied
by the sciences and the types of representations used by scientists” (Mitchell 2000,
p. 243). She defends a “multidimensional account” of scientific knowledge, proposing
a multi-dimensional conceptual space that spans the axes of abstraction, stability
and strength. This scheme allows for a more comprehensive conception of “law” in
which both the law of conservation of mass and Mendel’s law can be represented in
the same conceptual space, with the latter just being less stable. Thus the advantage
of the model is that “the strength of the determination can also vary from low prob-
ability relations to full-fledged determinism, from unique to multiple outcomes”
(Mitchell 2000, p. 263).
Once the multidimensionality of scientific knowledge is acknowledged, one
might even go further and admit that scientific knowledge is not only a matter of
concepts and theories but also a matter of practice. This is an issue raised by
Shreder-Frechette and McCoy when they argue that the “logic” of case studies
consists in a certain pragmatic procedure which ultimately makes them a reliable
basis for comparison as well as generalization. This is because the rationality of
practice relies on rules and on the reference to a community of individuals – just as
scientific concepts themselves do. “Hence the ‘logic’ of case studies may be
7 Waters’s oft-quoted example of law-like causal regularity is the following: “Blood vessels with
a high content of elastin expand as internal fluid pressure increases and contract as the pressure
decreases.” (Waters 1998, p. 19).
8 For further details, see Weber (1999) and Schweitzer (2000). Drawing on Waters’s analysis, Weber
eventually draws the not so surprising conclusion that “ecology knows evolutionarily invariant
generalizations which are law-like and at the same time distinctively biological.” (1999, p. 71).
126 A. Schwarz
appropriate to science if one conceives of scientific justification and objectivity in
terms of method, in terms of practices that are unbiased – rather than in terms
merely of a set of inferences, propositions that are impersonal” (Shrader-Frechette
and McCoy 1994, p. 243). Thus the uniqueness of case studies must be seen to lie
not only in their enactment of purely subjective rules and untestable principles.
Instead, ecological case studies should be perceived and appreciated epistemologi-
cally for the specific form of knowledge they manifest, which accommodates prac-
tical as well as conceptual and methodological analysis. It is likely that this
epistemic form is closer to the ideal of a proper epistemic model in ecology.
Thus there are a range of promising approaches available within the philosophy
of science to enable an adequate description of a corpus of knowledge as plural in
terms of its methods, practices, concepts and objects as that generated in scientific
ecology. The approaches discussed,9 be they Cooper’s concept of nomic force,
Carrier’s concept of supervenience, or Mitchell’s multidimensional space – all
these expounding a more differentiated gradation of scientific generalizations – or
again Shrader-Frechette and McCoy’s proposal of a “logic” of case studies based
on pragmatic procedures: all these approaches provide appropriate instruments for
describing ecology as a powerful scientific discipline.
A Rational Reconstruction of Ecology
The question now is how the conceptual triad grounded in natural philosophy can
be linked with the dynamics of theory development: How do the basic conceptions
and thus ecological theories take over where the others leave off, what is the con-
nection between the natural philosophical content – the “character” – of a basic
conception and the theories developed within each one? Which philosophy of sci-
ence is capable – to borrow a meta-history of science norm from Lakatos as a cri-
terion of quality – of integrating “history of science” the most, that is, of
reconstructing the most concepts and theories in a rational manner? The philosophy
of science method of rational reconstruction proposed by Lakatos certainly does
seem best suited to describe adequately the dynamic development of theory in ecol-
ogy. This is because, first, the method does not prescribe its methodological con-
struction according to criteria set by some external authority but rather orients itself
towards the judgment of the community itself regarding the extent to which a the-
ory is regarded as progressive or degenerative. This provides a good depiction of
ecological relationships, because it is precisely here that these latent simultaneities
of research programmes are encountered. Second, methodological strategies are
proposed in order to tie the dynamics of theory building back to a natural
philosophical substance, or “natural character”. An additional third argument
might be that Lakatos holds not only that “the history of science is the history of
9 To rule out the possibility of misunderstandings at this point: none of the approaches mentioned
lays claim to completeness.
1278 Dynamics in the Formation of Ecological Knowledge
research programmes rather than of theories” but beyond that this “may therefore
be seen as a partial vindication of the view that the history of science is the history
of conceptual frameworks or of scientific languages.” (Lakatos 1972, p. 132).
Research programmes consist of a “hard core” which is adhered to even when
difficulties with experiments arise, and a “protection belt”, in which auxiliary
hypotheses are developed and adjustments made in order to protect the core. The
adjustments are guided by a positive heuristic, which defines problems and makes
methods available. It is the “protective belt of auxiliary hypotheses which has to bear
the brunt of tests and get adjusted and re-adjusted, or even completely replaced, to
defend the thus-hardened core” (loc cit., p. 133). The heuristic power of a research
programme is the capacity to anticipate novel facts in its growth. Thus, a research
programme is successful as long as its progressive problemshift continues. “Creative
imagination is likely to find corroborating novel evidence even for the most ‘absurd’
programme, if the search has sufficient drive.” (loc cit., p. 187). Lakatos even admits
that the increase in content needs to prove its worth only occasionally in retrospect
in order to have enough scope for giving a rational explanation for dogmatically
clinging to a research programme (even in the face of refutations): how many new
facts did it produce, how great was the capacity of research programmes “to explain
their refutations in the course of their growth” (loc cit., p. 137).
This idea makes it possible to say, then, that the refutations are able to have no
impact on the hard core: Thus research programmes are scientific achievements
which can be evaluated on the basis of progressive and degenerating problemshifts.
The basic unit of evaluation for progressiveness or degeneration is not the individ-
ual theory10 but a series of theories – the research programme. The function of the
research programme consists in predicting anomalies through its positive explana-
tory potential and integrating them by force of the dynamics of persuasion. What is
decisive is that the research programmes are simultaneously characterised by a
sluggishness which guarantees their continuity. This continuity can not simply be
interrupted by a “crucial experiment” or an individual theory, either. Lakatos states
that in this sense no experiment can ever be regarded as decisive, neither at the time
it was conducted, and certainly not beforehand.
The replacement of one research programme by another occurs when it “loses
its empirical power”, proposes Lakatos by paraphrasing Popper (loc cit., p. 154). A new
programme already developed in nuce during the lifetime of the old programme
comes to compete with the older one and eventually outstrips it by its greater
problem-solving power. “While the old programme increasingly requires theoretical
adaptations to uncooperative experiential data (degenerating problem shifts), theoreti-
cal development in the new one maintains the upper hand (progressive problem
shift): empiricism becomes not the task of the new programme but a successful test
of it” (Diederich 1974, p. 14). A research programme continues to advance as long
10 Diederich (1974) and other authors (e.g. Wolfgang Stegmüller) complain that Lakatos’s use of
the term “theory” is not as unequivocal, as the powerful weighting of theory towards research
programmes would suggest with some justification.
128 A. Schwarz
as new facts can be predicted with a certain degree of success,
and it stagnates if its theoretical growth is retarded – if it is able to offer only post-
hoc explanations. The point at which it is completely cancelled is when a new
research programme is capable of demonstrating greater explanatory potential
compared with its predecessor.
However, what is key here is that Lakatos admits of competition between two
research programmes as a “long extended process, during which it is possible to
work rationally on each of the two programmes (or, if possible, on both)” (Lakatos
1974, p. 282; emphasis in original). The difficulty of deciding when a research
programme can finally be regarded as having been superseded could be made fur-
ther acute insofar as the extended process of competition between two research
programmes can not, in principle, be contained at all. “Even when a research pro-
gramme is seen to be swept away by its predecessor, it is not swept away by some
‘crucial’ experiment; and even if some crucial experiment is later called in doubt,
the new research programme cannot be stopped without a powerful progressive
upsurge of the old programme.” (Lakatos 1972, p. 163). Thus research programmes
would exist in perpetuity and – depending on the one hand on the internal develop-
ment of the degenerating problem shift and on the other on external societal
developments – alternately put the other “rival to one side”, which then lurks in the
background, awaiting its opportunity to reconquer the field. However, Lakatos
points out that there lies no necessity in the series of alternation of speculative
conjectures and empirical refutations. Rather, the “dialectic of research pro-
grammes” is characterized by a broad variety: “which pattern is actually realized
depends only on historical accident.” (loc. cit., p. 151).
This “dialectic” and the variety of interactions between the evolution of the pro-
gramme and the empirical checks as well as the simultaneity of co-existing research
programmes seems to be a fitting description of the plural situation in ecology. This
is the sense in which the three ecological basic conceptions of “microcosm”, “energy”
and “niche” will be discussed in the following. In the descriptive mode of the
Lakatosian research programme, the basic conceptions’s natural philosophical con-
tent (or natural character) merges and becomes a part of the hard core.
Three Basic Conceptions in Ecology
The purpose of the term “basic conception” is to describe the possibility of three
conceptions of nature and to render them operational in such a way that, by provid-
ing a normative pattern, they in turn render the discipline of ecology comprehen-
sible. This pattern opens up and simultaneously limits the space of possibilities for
concept formation and theory building. The partial knowledge areas and the narratives
present in ecology are structured along these lines. The basic conception of microcosm,
for example, typically contains a romantic narrative, whereas the basic conception
of “niche” is characterized more by a drive economic narrative based on the
economy of drives. This pattern of basic conceptions turns out to be quite robust;
in a certain sense it might even be called archaic: it is linked to the very beginnings
1298 Dynamics in the Formation of Ecological Knowledge
of ecology as a scientific discipline, has been reproduced time and again ever since,
and serves as a rather intuitive meta-narrative of the discipline. While the triadic
pattern is systematically robust, it is historically flexible insofar as the power of
representation of the basic conceptions among each other can be switched and
repositioned, depending on which character of nature fits best with the narrative or
heuristics concerned and eventually helps to get a research programme accepted.
Which of the three conceptions of nature is able to become established in a specific
historical situation is subject not only to conditions within science itself but is also
influenced by societal projections and expectations of ecological concepts or mod-
els. In order to get an impression of the success, historically speaking, of a particu-
lar basic conception, and to assess the joint oscillation of all three basic conceptions
over a longer period of time, it is important not only to pay heed to the dynamics
between the three characters of nature and those of the formation of concepts and
hypotheses, but also to consider which socio-political expectations and prevailing
circumstances11 the ecological description of nature refers to and builds upon (and,
conversely, which ones it influences).
Basic Conception of “Niche”
Within the basic conception of “niche” organisms are imagined as “self-determined”
individuals in early ecology. In “the lake as a microcosm”, a paper that was influ-
ential in the historical development of ecology, American zoologist Stephen Alfred
Forbes (1887) describes organisms as “remarkably isolated” and indifferent or even
hostile towards one another. There is an abundantly clear reference in Forbes to a
economic driven character of nature and therefore to “niche”, and this is confirmed
by closer analysis at the conceptual level.12 Predator-prey relationships are included
in the field of view here; they appear to be ubiquitous and the relationships between
the competing individuals are influenced by the principle that every animal has its
enemies, and “mercy and welfare are completely unknown” (Forbes 1887). These
organisms come together in societies in which the distribution of roles are deter-
mined solely according to the purpose of the association: the organisms build a
society based on what might be called a “social contract”. The organisms’
11 Particularly Ludwig Trepl and collaborators underline the socio-political influence on ecological
theories; beyond this rather general claim they see a close relationship between a distinct political
constellation and the concepts of ecological society or community (see Trepl 1987, 1994, and also
Trepl & Voigt Chap. 5, this volume).
12 It may at first seem confusing that the very publication in which the lake is proclaimed – for the
first time – to be a microcosm is not enlisted for the basic conception of “microcosm”. Here,
though, microcosm refers above all to the idea of a system per se, a combined view of individual
parts which at that point could not yet be integrated conceptually. But this new perspective is in
some sense the ecosystem perspective and can occur in all three basic conceptions. So the phrase
“lake as microcosm” is not on its own sufficient for deciding to which of the basic conceptions it
should be ascribed.
130 A. Schwarz
environment – biotic and abiotic – is at most a neutral, but more usually hostile
affair, which intervenes in the life of the individuals for purposes of regulation.
The Darwinian principle of natural selection is omnipresent here, so that the
evolutionary biological model of the niche is able to become quite well established
as a result. One of the most successful ecological theories is developed in this con-
text, namely the “competitive exclusion principle“ (CEP). The CEP states that two
species coexisting in the same space compete for limited resources. The niche is
regarded as a spatially organized structural unit which the competing species endea-
vour to gain possession of. The structure of the niche lies to a certain extent at the
bottom of the society, whose stability is an outcome of the regulation that occurs
through competition between the species. The CEP is held to be a model which is
so well developed mathematically that it is also considered capable of making pre-
dictions. It is conceived as a key to explaining the composition of animal and plant
societies, leading to the dominance of the basic conception of “niche” in theory
building in ecology (Gause 1934).
When Charles Elton (1927) introduced the functional concept of niche, the CEP
was temporarily in trouble. What Elton wants to achieve with his conception of
niche is to describe the position of an animal in the food chain and therefore its
economic significance. This new function-based niche did not entirely supplant the
old spatially-conceived niche. The old meaning of niche as habitat and not as a posi-
tion in the food chain crops up at various points (Kingsland 1991, p. 6). So the CEP
also plays a role in Elton’s conception, albeit not a dominant one; the principles of
competition and selection receded into the background in Elton’s work. Ultimately,
the heuristic of the spatial concept of niche becomes weakened as a result. In Elton’s
work the concept of niche is transformed: the spatial niche becomes the functional
niche and at the same time is “transferred from the evolutionary biological context
into an ecological context” (Trepl 1987, p. 170). This “ecological context” fits with
the basic conception of “energy” in the triadic conception, where relational condi-
tions and a systemic perspective play a role above all.
However, this is by no means the end of the evolutionary biological concept of
niche: it experiences a renaissance in the 1950s, especially in plant ecology, and is
then brought successfully to bear in opposition to the so-called climax theory (basic
conception “microcosm”). Eventually, the functional and the spatial concept of niche
are integrated in a single theory: the niche concept is conceptualized as a geometric,
“n-dimensional hyperspace” (Hutchinson 1957) which makes it possible to depict all
ecological factors, i.e. the sum total of a species’ conditions of existence – upon
which the building of successful theory takes place in the context of the basic concep-
tion of “energy”.
Basic Conception of “Microcosm”
The “microcosm” relates to romantic nature. As a more or less well thought-through
position in terms of philosophy of nature, this nature can currently be found, for
example, in nature conservation ecology, but is also quite a widespread narrative in
1318 Dynamics in the Formation of Ecological Knowledge
theoretical ecology. This applies all the more to early ecology at the close of the
nineteenth century. Descriptions of a lake as the “arena of life”13 or as the “self-
enclosed, clearly demarcated whole” (Forel 1901), or again as a “mirror image of
processes in the larger whole” (Zacharias 1905) in which an “extremely intricate
living hustle and bustle” (ibid.) is found, are not unusual. The principle of selection
at the centre of the basic conception of “niche”, that of the control of individual
organisms from the outside, is replaced in the basic conception of the “microcosm”
by principles of reciprocal and (usually) highly hierarchical control. As a result, the
biological community is viewed not according to the premise of competition
between individuals but according to the premise of fitting in or adaptation to the
community, along with an emphasis on the interplay and the mutual dependency
among all individuals or species. Here, the local conditions are not an expression of
relentless natural laws by which the plants and animals occurring more or less by
chance in a certain place are externally controlled. Instead, the community actively
adapts to the local conditions, and in the process of adaptation, these local conditions
are “internalized”.
In plant ecology, for example, this position has been represented by the “super-
organism” theory, in which the plant community is conceptualized as a super-
organismic individual, whose individual organisms exist in intra-societally organized
functional interrelationships. The superorganism successively works to develop its
individuality through the interrelations between organisms and general local condi-
tions, which at the same time are a specific place. In this process of adaptation the
plant community increasingly detaches itself from the immediate and specific con-
straint of nature and simultaneously adapts to the region and climate. The total
organism “plant community” develops the specific aspects of the place, which dis-
tinguish it from all other places and the organisms adapted to them. Since the plant
community contributes to shaping the space surrounding it, though, its relationship
to this space is not exclusively one of dependence but rather highlights the particu-
larity of the organized individual.
Basic Conception of “Energy”
The introduction of the conception of energy in physics established a unity between
different sub-fields of physics.14 Qualitatively different natural phenomena can not
only be “reduced to a common one in the conception of energy”; more than this,
13 Forbes 1887; Forel 1891, 1901; Zacharias 1904, 1905, 1907, 1909.
14 “Unwittingly perhaps, Helmholtz was the first great bourgois philosopher of labor power pre-
cisely because in his essays on Kraft he does not distinguish between natural, mechanical, or human
labor power. For him all expenditure of energy produced work, and conversely all work involved
the consumption of energy. His conception of labor power reveals no self-moving power, no social
labor that is not at the same time a natural force. With Helmholtz work was reduced to a quantitative
phenomenon subject to a system of mathematical equivalents” (Rabinbach 1990, p. 61).
132 A. Schwarz
“energy is seen in some sense as the origin of all changes in nature and as the measure
of all impacts” (Breger 1982, p. 41). It just seemed to be a matter of consistency
that, after the conception of energy had been introduced in physics, within a short
time it became common practice to describe all natural phenomena by giving their
energy distribution.
In the physiology of the 1850s this idea of the standardization of the forces of
nature using the concept of energy was taken up in something of a vulgar material-
istic variation. The “eternal cycle of material-bound life”, in fact, life per se, was
declared to be “nothing but material”, which in turn was supposed to be nothing
other than force or energy.15 A material characterized in this way can quite easily
be non-living or living nature, and could now be conveyed on a material basis via
the cycle.
This is taken up in early aquatic ecology, for instance with the notion of the
circulating “organic substance” in a lake. This organic substance was not character-
ized by the property of a particular material or of a chemical element but by its
function of connecting the organisms in the lake with one another and with their
inorganic environment. Organic substances are materials with which work is done
on an organism, i.e. ones which contribute to its maintenance or are brought forth
by it. The crucial point is that these functions of organic substance are always
related to the “lake as a system”, in which the cycle of organic substance takes
place. By analogy to the physical conception of energy, the reference here, too, is
to an image of nature in which the idea of balance and of unity are of central sig-
nificance – we are looking at a nature of control.
Provisional Result
The simultaneity of co-existing research programmes seems to be a fitting descrip- –
tion of the plural situation in ecology. This is the sense in which the three ecologi-
cal basic conceptions of “microcosm”, “energy” and “niche” were discussed.
The three basic conceptions provide a normative pattern that opens up and –
simultaneously limits the space of possibilities for concept formation and theory
building. Concepts and narratives present in ecology are structured along these
lines.
This pattern of basic conceptions turns out to be quite robust; it has been repro- –
duced time and again ever since the scientific field of ecological knowledge
exists, and seems to serve as a meta-narrative of the discipline.
15 In physics the concept “energy” is shaped not before the 1850s. However, a unifying force that
goes far beyond Newton’s concept of force, is already in discussion much earlier (see for instance
Schelling) (Breger 1882, p. 98 f.).
1338 Dynamics in the Formation of Ecological Knowledge
The triadic pattern is flexible over time in so far as the power of representation –
of the basic conceptions among each other can be switched and repositioned,
depending on which character of nature fits best with the narrative or heuristics
concerned and eventually helps to get a research programme accepted.
Which of the three basic conceptions is able to become established in a specific –
historical situation is subject not only to conditions within science itself but is
also influenced by societal projections and expectations of ecological concepts
or models.
Oscillating Basic Conceptions
A crucial part of the theoretical dynamics described here is that the basic concep-
tions are able to coexist. Which of the basic conceptions is able to predominate over
the others seems to be less a question of the internal theoretical dynamics and more
one of the degree of fit between the structural core, or character, of the basic con-
ception and the socio-political context.
In early ecology it was the basic conception of “niche” that came powerfully to
the fore.16 Towards the end of the nineteenth century plant ecologist Eugen
Warming (1841–1924) emphasized that competition should be regarded as the most
important relationship between the members of a plant community, thus aligning
himself with a prevailing “Darwinian paradigm”. “Community” was conceived of
as a collection of individuals that confront each other in the competition over scarce
resources in either an indifferent or a hostile way. From this perspective, individuals
and species in a given location live in a biological community, but are largely deter-
mined by their individual biological capacities. In liberal social scientific theories,
the contradiction between the active component in this model – the active individ-
ual fighting for survival – and the passive component – the external exigencies of the
environment – is resolved in an endless and undirected evolutionary process, that
is, in progress. This heavy emphasis on endlessness and progress in Darwinian
theory and later in Gleason’s theory of succession (Gleason 1926) became a point
of widespread criticism in ecological community, ringing in – as historian and theo-
rist of ecology Ludwig Trepl argues, for example – a conservative phase. From the
conservative perspective the biological community does not function on the premise
of competition between individuals but on the premise of adaptation and an emphasis
on the interplay and the mutual dependency among all individuals or species. By
analogy to the conservative social philosophy, this construction corresponds to the
idea of the “superorganism”: the whole comes before the parts, which fulfil their
task in a functionalist, teleological way. The “superorganism” contains and combines
16 This was discussed in more detail in Schwarz and Trepl (1998, p. 305 f ).
134 A. Schwarz
the cooperating elements into a harmonious structure of a higher order (Clements
1936, but also already 1916). The conservative idea of the state is also characterized
by the endeavour to eliminate the liberal distinction between state and society or at
least to weaken it, so that the community (which ultimately means an indivisible
unity of state and society) acquires a more important role than the one assigned it
by the liberal position:
The societal organism which is recognized as relatively higher must be not only that in
which greater scope is given to the political freedom of the individual citizens or social
groups, but also that in which state power in its greatest concentration also possesses the
greatest independence, self-determination, and freedom of action. (Lilienfeld 1873, p. 84)
In ecological theories of the romantic-holistic type, similar structures which arose
at roughly the same time can be identified:
Every oyster bank is in some sense a community of living creatures, a selection of species
and a sum of individuals which find in this very place all the conditions for their emergence
and maintenance, that is, the right soil, sufficient food, the appropriate salt content and
tolerable temperatures favourable for development. … All living members of a living com-
munity keep […] the balance with the physical conditions of their biocoenosis because they
maintain themselves and reproduce themselves in the face of all the influences of external
irritations and against all attacks on the continued existence of their individuality. (Möbius
1986 (1877), p. 74 f.)
In the 1920s and 30s the idea of superindividual units became dominant in aquatic
ecology as well. Richard Woltereck (1877–1944) concerned himself extensively
with the “spatial structures of biosystems” (title of chapter eight; Woltereck 1940
(1932), p. 208) and developed an intricate terminology in which individuals were
conceived of as “components of multi-sectional systems (collective structure)”. The
so-called “multi-person systems” were able to be “physically unified edifices”
which “belong together only in spatial terms”. Edifices are structures used to
describe an “ordered multiplicity”, and between whose “components interactions
take place which are merged together by superordinate relationships into a whole
or gestalt. This applies to the individual organism as a self- or individual edifice
[…], secondly to the organism in its environment […], thirdly to multi-sectional,
but self-enclosed systems of organisms” (ibid.). All “three kinds of system” are to
be conceived of as “edifices of relationships”.
Woltereck distances himself vigorously from the “niche position” and thus simul-
taneously from political liberalism: Chance, according to Woltereck, was “nothing
but a shabby word, a non-concept and a nonsense (…), despite the supposedly cre-
ative power which the term ‘selection’ was to infuse into this chance.” (l.c., p. 230).
In the years after 1930 this holistic image of nature as “romantic nature” was
gradually replaced by the basic conception of “energy” and thus by a synthetic
position between so-called reductionism and organicism (or vitalism). However,
Ludwig von Bertalanffy writes already in 1929 (p. 95):
We understand on the basis of the maintenance of gestalts how the organism preserves
itself in metabolism. …Our view is a theory of a system remaining steadfast in its condition
and overcomes both machine theory, which is not adequate, as well as the scientifically
impossible vitalist view.
1358 Dynamics in the Formation of Ecological Knowledge
This position became highly successful in ecology; the early ecosystem theories
were developed out of the basic conception of “energy” (Schwarz 1996). Ecosystem
wholeness promised controllability and the technical reconstruction of nature.
What Holds Ecology Together?
The question remains, however, why a plural ecology does not fall apart but instead
remains a reference system to which research programmes and concepts are bound
and a particular corpus of knowledge refers, even when the disciplinary edges
become blurred. Why does a system – be it an experimental system,17 a discipline,
or a research programme – peters out (or not)?
A variety of epistemological proposals have been presented, each referring to
different systemic levels: The experimental system becomes exhausted when there
are too many irreconcilable, conflicting answers available; the heuristic of a
research programme becomes negative when its theories are no longer able to
explain the phenomena observed; and a discipline falls apart when its instruments
and methods become progressively diverse while other epistemic objects, or “ques-
tion machines” (Rheinberger 1992, p. 72), emerge.
The approach proposed in the following is based above all on an analysis of
concepts as constitutive elements of ecology as a knowledge system. It therefore
argues that it is mainly the conceptual architecture, rather than theories, methods or
instruments that guarantees disciplinary and logical robustness. This does not
mean, of course, that in the triadic design of the basic conceptions of energy, niche
and microcosm theories or narratives are not equally at work. After all, concepts
only function as such when they are appropriately contextualized – in distinct theo-
ries or narratives. Thus there is typically a romantic narrative contained in the basic
conception of microcosm, for example. The basic conception “niche” tends to be
influenced by an economy of drives, while the conception “energy” relies on nar-
ratives in which functional und systemic relations play a prominent role. Based on
the triadic approach, then, the temporal and epistemic stability of ecology can be
construed as follows:
It is generated (a) by the interrrelational context in which the concepts are
embedded, represented by distinct theories and narratives; (b) by the dynamic
which links them, and finally (c) by being tied to the historical genesis of a triadic
concept of nature.
In order to strengthen the latter claim, a philosophy of history approach is
mobilized which is aimed at a pluralistic concept of nature. This is where the tri-
adic construction of the basic conceptions finds a philosophical rationale.
17 According to a conception developed by Hagner et al. an “experimental system” in laboratory sci-
ence is one in which differences are generated (Hagner, Rheinberger, Wahrig-Schmidt 1994, p. 10).
136 A. Schwarz
Historical Genesis a Rationale for Pluralism in Ecology
Based on the History of Philosophy
Based on the above, three “characters of nature” can be ascribed to the three eco-
logical basic conceptions. Working on the basis of Marquard’s ideas, the status of
these characters of nature is important in systematic terms. The basic conception of
“niche” is supported by drive nature, that of “microcosm” by romantic nature, and
the basic conception of “energy” is closest to controlling nature.
The historical genesis of the ecological triad can be localized around the turn of
the nineteenth century. In the first instance this is not especially surprising because
it coincides with the general perception of a historical break at that time, particu-
larly for the sciences of biology and chemistry. A myriad of studies has been pub-
lished probing various issues in this regard; however, most of them share the
following story: The relationship between nature and culture changes more or less
radically, this being apparent in the reorganisation of conceptual frameworks in
philosophy, science and literature, in the situatedness of ontology, and also in the
narrative forms. These changes permeate through to all scientific disciplines and
societal discourses and practices in equal measure. The accelerated secularisation
of the order of Nature is reflected in a gradual shift of traits from the outside of
bodies to inside them, from concrete and visible signs to abstract and unseen ones.
A mathematically describable and law-like Nature steps up to compete with the
perception and representation of Nature as a hierarchical chain of beings – the vis-
ible signs of the hand of God (allegorically speaking). At the “end of natural his-
tory” (Lepenies 1976, Latour 2005) we are left with a Nature which is split off from
everything, which is considered to be culturally conditioned – a split which is also
reflected in the distinction between two concepts of history: the history of Nature
and the history of humans.18 Historians and philosophers place temporal and spatial
limits on this historical break have it lasting for a longer or shorter time, and con-
sider it to be more rhetorically than pragmatically effective – or vice versa. Usually,
however, it is identified with the beginnings of modernity and of a construction of
objectivity in scientific knowledge production, which relies on what Feyerabend
has called the “separability thesis”.
In environmental philosophy, debates abound concerning the shift that occurred
in the concept of Nature at that time and also concerning the traces of these con-
cepts in contemporary theories and practices of perceiving and representing
Nature. Much less common is the effort to link these philosophical conceptions
together in order to substantiate the epistemology of ecology, as proposed in the
following.
18 In Schelling the latter is merely “inauthentic” history which, moreover, is inferior to the authen-
tic history of nature, a history of the whole of Nature (natura naturans), of Nature that is constantly
reproducing itself. This conception of a processual Nature is an oft-used figure of thought in
environmental philosophy. (For a review of Schelling’s work in the field of ecocriticism, espe-
cially with respect to wilderness, see Wilke 2008 from a perspective of literary criticism).
1378 Dynamics in the Formation of Ecological Knowledge
Philosopher Odo Marquard conceptualizes the historical break in which the
hitherto fixed relationships between reason, humanity and nature have to be refor-
mulated as a period of transition. The modern constitution of nature as “the sphere
opposite to reason” (das Andere der Vernunft) (Böhme and Böhme 1983) is not yet
consolidated conceptually in this historical phase, manifesting in a “triadisation” of
the concept of nature. The conceptual triad is a snapshot of relationships that
remain blurred, within which the new relations emerge.
The thesis put forward in this paper, then, is that these three concepts of nature
and their relations and references to and among one another contribute towards
stabilising the plural local and temporary development of concepts and theories in
ecology, up to and including the present day. Latour’s “we have never been modern”
applies to ecology to the extent that the latter never obeyed the dominant narrative
of a “work of purification” identified with modernity, either historically or system-
atically. This narrative is replaced in the triadisation of the nature concept by a
constellation in permanent transition.
A Triadic Conceptualization of a Philosophically
Delocalized Nature
Marquard goes along with the common mode of reading which sees nature, when
the historical break19 occurs, become the estranged and split off counterpart, the
other of reason. At that moment, the concept of nature is no longer bound up as part
of a metaphysically conceptualized world; instead, it is constructed through scien-
tific experience, which is oriented towards control and imagines nature as a subject
to making and remaking (“machbare Natur”). As a result, statements about the
essence of nature become “dispensable, or rather impossible”.20 This constellation
gives rise to a dilemma, which leads to a shift in the concept of nature along with
a process of differentiation. The philosophical reformulation of the concept of
nature is challenged, according to Marquard, by a reason that has “begun to waver”.
Reason threatens to depart from human nature – something that cannot actually
happen, as the very realization of human nature itself is identified with reason.21
19 On the question of a “reasonable” construction of an epochal break see Nordmann (2006).
20 Mittelstraß 1981, p. 64. In philosophy the historical break is characterised by the transition from
transcendental philosophy to philosophy of history. This proves to be a dilemma in the context of
the “anthropological reorientation towards nature” regarding the “question of essence”, which in
the context of the philosophy of history can only be a false question: the transcendental philo-
sophical framework has been dissolved.
21 In Marquard’s words, human nature is “tied to reason and to that which makes reason what it is,
what enables it to flourish” (Marquard 1987, p. 54). Drawing on the Aristotelian concept of nature,
M. describes this as “actual” nature. If we attempt to subtract human reason from human nature, it
will continue to strive “in itself” towards a realization of reason, without being able to find it – a state
which, as “potential nature”, is in opposition to actual nature. To assert that this potential nature is
one that is completely emancipated from reason is to accept that it is a “mere possibility”.
138 A. Schwarz
Philosophically, he argues, nature in this situation can no longer be conceptualized
within a transcendental context and cannot yet be conceptualized through the phi-
losophy of history. This situation of a delocalized nature explained by Marquard
through reference to the interplay of three concepts of nature: controlling nature
(Kontrollnatur), romantic nature (Romantiknatur) and drive nature (Triebnatur).
Drive nature refers to lack of control, to the satisfaction of individual desires,
and to the realm of the senses in general. Hobbes is called as the crown witness in
this context, with his famous dictum: “Hereby it is manifest, that during the time
men live without a common Power to keep them all in awe, they are in that condi-
tion which is called Warre; and such warre, as is of every man, against every man”
(Hobbes 1651, p. 62). When there is no common, binding set of rules to channel
passions, the result, says Hobbes, is a civil war in which everyone fights against
everyone else. Only fear in the face of a superordinate power can keep humans from
hurtling headlong into violence and lack of restraint. This natural state of human
beings – of humanity per se – is ever present, even if it is not always equally power-
ful. It gives a society (though not a community), a particular character that is
opposed to the state: “drive nature is present (in tempered form) as society”
(Marquard 1987, p. 55).
Romantic nature is above all nature perceived as organism; it can be exalted and
beautiful. Whether manifesting as landscape or as fertile wilderness, the “condition
of innocence and naivety” (Marquard 1987, p. 57) is ever present in it. Contained
in romantic nature are notions of a harmonious cosmos, a symmetrical relationship
between microcosm and macrocosm, as well as that of a complete, whole primeval
state. Nature conceptualised as romantic also refers to inwardness and tempera-
ment, to feeling and longing, to the realms of imagination in general. As a conse-
quence it is especially the domain of literary fiction. “The poet understands nature
better than the scientist” (Novalis, quoted in Marquard 1987, p. 57). Only in the
presence of the poet does it show itself and reveal its secrets. This is why – in
Novalis’s interesting reversal of the argument – genuine “physics [is] nothing other
than the study of the imagination”: “the more poetic, the more true”. Romantic
nature is above all nature perceived and interpreted aesthetically.
Finally, Marquard identifies controlling nature as that of the exact sciences. It is
that which, through observation, experiment and mathematics, can be recorded,
manipulated, predicted and therefore controlled. Above all, however, this nature is
not “merely sensual” nature but the nature of rational rules, the nature of the ratio-
nal mind. It is not only derived from the laws of the possibility of experience: it is
identical with them. For Marquard, Kant is the source for controlling nature with
his famous statement that the general laws of nature must be settled independently
(that is before every experience). According to Marquard then, controlling nature is
the nature of science and technology, present as “design, method, result and appli-
cation” (1987, p. 56).
Thus only the last of these three concepts of nature, controlling nature, is one
relevant to the natural sciences. This constitutes a pronounced difference in com-
parison with the ecological basic conceptions, all three of which aim to provide an
opening into science. What Marquard’s scheme certainly does do, however,
is provide a historical rationale for the pluralization of the concept of nature and a
1398 Dynamics in the Formation of Ecological Knowledge
philosophical rationale for the triadisation. Both are useful here in order to fashion
a systematically sound foundation for the triadic conceptualization of ecology. In
this sense delocalized nature – that is, the triadization of the concept of nature – is
viewed as a scheme rooted in natural philosophy, which underlie the basic concep-
tions, acting as a framework which, from a philosophy of science perspective, can-
not be circumvented. Unlike hypotheses, such frameworks are exempted from
critical examination, being used instead to guide the activity and orientation of
hypothesis formation.
This interconnection between hypothesis formation exposed to empirical verifi-
cation protected from it has been described in the context of various approaches in
the philosophy of science. The “metaphysical core” in Imre Lakatos’s research
programmes22 as already discussed and the “themes” in scientific thoughts as
described by Gerald Holton are examples. Similarly, Gernot Böhme proposes the
notion of physiognomically deducible “characters”, namely “characters of nature”
(Naturcharaktere) which are at work behind the epistemic operations ascribing
meaning to nature. Characters of nature are
sentences in which certain character traits are ascribed to nature, [they] are an expression
of highly aggregated experiences. They name an overall impression of nature that is
grasped intuitively and formulate it in physiognomic manner: As nature shows certain
basic traits, it is ascribed a character, as it were; it is said what is basically expected from
it. Statements about the character traits of nature are made not on the basis of scientific
experiences alone. For science, they perform the function of heuristics, that is, of instruc-
tions for what should be sought in nature and what is to be expected of it. Statements about
characters of nature are thus still related to science and its transformation, but they are not
scientific statements per se. They can be understood as genuine statements of a philosophy
of nature to the extent that the latter is an endeavour to understand nature as a whole and
as such. (Böhme 1992, p. 211)
In this sense the three basic conceptions provide a theoretical foundation for ecol-
ogy based on the philosophy of nature: they make explicit three conceptions of
nature which are identified historically with the beginning of modernity and are
capable systematically of stabilizing the plural representation of this science.
References
Allen TFA, Tainter JA, Chris Pires J, Hoekstra T (2001) Dragnet ecology “Just the facts Ma’m”:
The privilege of science in a postmodern world. Bioscience 51:475–484
Beatty J (1995) The evolutionary contingency thesis. In: Wolters G, Lennox JG, McLaughlin P
(eds) Concepts, theories, and rationality in biological sciences. The Second Pittsburgh-
Konstanz Colloquium in the Philosophy of Science. University of Pittsburgh Press, Pittsburgh,
pp 45–81
22 It is important to note here that Lakatos makes no difference between the methodology of a
research programme with a “metaphysical” core and “one with a ‘refutable’ core except perhaps
for the logical level of the inconsistencies which are the driving force of the programme.” (Lakatos
1972, p. 127).
140 A. Schwarz
Böhme G (1992) Wissenschaft-Technik-Gesellschaft. 10 Semester interdisziplinäres Kolloquium
an der THD. TH Darmstadt, Darmstadt
Böhme H, Böhme G (1983) Das Andere der Vernunft: zur Entwicklung von Rationalitätsstrukturen
am Beispiel Kants. Suhrkamp, Frankfurt/M
Breger H (1982) Die Natur als arbeitende Maschine. Zur Entstehung des Energiebegriffs in der
Physik 1840–1850. Campus, Frankfurt/M
Carnap R (1956) Meaning and necessity: a study in semantics and modal logic. (Reprinted of
Revue Internationale de Philosophie 4 (1950): 20-40). University of Chicago Press, Chicago
Carrier M (1995) Evolutionary change and lawlikeness. Beatty on biological generalizations. In:
Wolters G, Lennox JG (eds) Concepts, theories, and rationality in the biological sciences.
Universitätsverlag Konstanz/University of Pittsburgh Press, Konstanz/Pittsburgh
Cartwright N (1999) The dappled world. A study of the boundaries of science. Cambridge
University Press, Cambridge
Clements FE (1936) Nature and structure of climax. J Ecol 24:252–284
Cooper GJ (1996) Theoretical modeling and biological laws. Philos Sci 63:28–35
Cooper GJ (1998) Generalizations in ecology: a philosophical taxonomy. Biol Philos
13:555–586
Diederich W (ed) (1974) Theorien der Wissenschaftsgeschichte. Beiträge zur diachronischen
Wissenschaftstheorie. Suhrkamp, Frankfurt/M
Elton C (1927) Animal ecology. Sidgwick & Jackson, London
Forbes SA (1887) The lake as a microcosm. Bull Peoria Sci Assoc 111:77–87. (Reprinted Bull
Nat Hist Surv 15:537-550, Nov 1925)
Forel F-A (1891) Allgemeine Biologie eines Süßwassersees. In: Zacharias O (ed) Die Tier- und
Pflanzenwelt des Süßwassers. Weber, Leipzig, pp 1–26
Forel F-A (1901) Handbuch der Seenkunde. Allgemeine Limnologie. Engelhorn, Stuttgart
Gause GF (1934) The struggle for existence. Williams and Wilkins, Baltimore
Giere R (1995) The skeptical perspective: science without laws of nature. In: Weinert F (ed) Laws
of nature. Essays on the philosophical, scientific and historical dimension. de Gruyter, Berlin/
New York
Gleason HA (1926) The individualistic concept of the plant associations. Bull Torrey Bot
Club 53:7–26
Haila Y, Taylor P (2001) The philosophical dullness of classical ecology, and a Levinsian alterna-
tive. Biology and Philosophy 16:93–102
Hagner M, Rheinberger H-J, Wahrig-Schmidt B (1994) Objekte, Differenzen, Konjunkturen. In:
Hagner M, Rheinberger H-J, Wahrig-Schmidt B (eds) Objekte-Differenzen-Konjunkturen:
Experimentalsysteme im historischen Kontext. Akademie Verlag, Berlin, pp 7–22
Hampe M (2000) Gesetz, Natur, Geltung. Historische Anmerkungen. Philos Nat 37:241–254
Hobbes T (1651) Leviathan, or, The matter, forme, and power of a common wealth, ecclesiasticall
and civil. Printed for Andrew Crooke, London
Hutchinson GE (1957) Concluding remarks. Population studies: Animal ecology and Demography.
Cold Spring Harbor Symposia on Quantitative Biology. T. b. L. C. S. Harbor. New York,
Long Island Biological Association. 22:415–422
Kiester RA (1980) Natural kinds, natural history and ecology. Synthese 43:331–342
Kingsland SE (1991) Defining ecology as a science. In: Real L, Brown JH (eds) Foundations of
Ecology. The University of Chicago Press, Chicago/London, pp 1–13
Lakatos I (1972) Falsification and the methodology of scientific research programmes. In: Lakatos
I, Musgrave A (eds) Criticism and the growth of knowledge. Cambridge University Press,
Cambridge
Lakatos I (1974) Die Geschichte der Wissenschaft und ihre rationalen Rekonstruktionen. In: Lakatos
I, Musgrave A (eds) Kritik und Erkenntnisfortschritt. Vieweg, Braunschweig, pp 271–312
Latour B (2005) From Realpolitik to Dingpolitik or how to make things public. In: Latour B,
Weibel P (eds) Making things public. Atmospheres of democracy. MIT Press, Cambridge,
pp 14–43
Lepenies W (1976) Das Ende der Naturgeschichte. Wandel kultureller Selbstverständlichkeiten in
den Wissenschaften des 18. und 19. Jahrhunderts. Hanser, München
1418 Dynamics in the Formation of Ecological Knowledge
Lilienfeld Pv (1873) Gedanken über die Socialwissenschaft der Zukunft. Behre, Mitau
Longino HE (2006) Theoretical pluralism and the scientific study of behaviour. In: Kellert SH,
Longino HE, Kenneth C (eds) Scientific pluralism. Minnesota studies in the philosophy of
Science. University of Minnesota Press, Minneapolis/London, pp 102–131
Marquard O (1987) Transzendentaler Idealismus, Romantische Naturphilosophie, Psychoanalyse.
Verlag Jürgen Dinter, Köln
McIntosh RP (1987) Pluralism in ecology. Annu Rev Ecol Syst 18:321–341
Mitchell S (2000) Dimensions of scientific law. Philos Sci 67:242–265
Mittelstraß J (1981) Das Wirken der Natur. In: Rapp F (ed) Naturverständnis und
Naturbeherrschung: philosophiegeschichtliche Entwicklung und gegenwärtiger Kontext.
Wilhelm Fink, München, pp 36–69
Möbius KA (2006) Zum Biozönose-Begriff. Die Auster und die Austernwirtschaft 1877 (2nd ed.
by T Potthast; 1st edition and comment by G Leps 1986). Harri Deutsch, Frankfurt/M
Motterlini M (2001) Reconstructing Lakatos. A reassessment of Lakatos’ philosophical project
and debates with Feyerabend in light of the Lakatos archive. London school of economics and
political science. Centre for philosophy of natural and social science. Discussion paper series
LSE (DP 56/01), pp 1–48
Nordmann A (2006) Collapse of distance: epistemic strategies of science and technoscience. Dan
Yearb Philos 41:7–34
Peters RH (1991) A critique for ecology. Cambridge University Press, Cambridge
Rabinbach A (1990) The human motor. Energy, fatigue, and the origins of modernity. University
of California Press, Berkeley
Rheinberger H-J (1992) Experiment, Differenz, Schrift: zur Geschichte epistemischer Dinge.
Basilisken-Presse, Marburg/L
Roughgarden J (1984) Competition and theory in community ecology. In: Salt G (ed) Ecology and
evolutionary biology: a round table on research. University of Chicago Press, Chicago
Ruse M (1973) Philosophy of biology. Hutchinson, London
Schwarz AE (1996) Gestalten werden Systeme: Frühe Systemtheorie in der Ökologie. In: Mathes K,
Breckling B, Ekschmidt K (eds) Systemtheorie in der Ökologie. Ecomed, Landsberg, pp 35–45
Schwarz AE, Trepl L (1998) The relativity of orientors: interdependence of potential goal func-
tions and political and social developments. In: Leupelt M, Müller F (eds) Eco targets, goal
functions, and orientors. Springer, Berlin, pp 298–311
Schweitzer B (2000) Naturgesetze in der Biologie. Philos Nat 37:367–374
Shrader-Frechette K, McCoy ED (1994) Applied ecology and the logic of case studies. Philos Sci
61:228–249
Smart JJC (1963) Philosophy and scientific realism. Routledge and Kegan Paul, London
Trepl L (1994) Holism and reductionism in ecology: technical, political, and ideological implications.
CNS 5:13–40
Trepl L (1987) Geschichte der Ökologie. Athenäum, Frankfurt/M
Waters CK (1998) Causal regularities in the biological world of contingent distributions. Biology
and Philosophy 13:5–36
Weber M (1999) The aim and structure of ecological theory. Philosophy of Science 66:71–93
Wiemken V, Boller T (2002) Ectomycorrhiza: gene expression, metabolism and the wood-wide
web. Curr Opin Plant Biol 5:355–361
Wilke S (2008) From ‘natura naturata’ to ‘natura naturans’: ‘Naturphilosophie’ and the concept of
a performing nature. Interculture 4:1–23
Woltereck R (1940) Ontologie des Lebendigen. Stuttgart, Ferdinand Enke
Zacharias O (1904) Skizze eines Spezial-Programms für Fischereiwissenschaftliche Forschungen.
Fischerei-Zeitung 7:112–115
Zacharias O (1905) Über die systematische Durchforschung der Binnengewässer und ihre
Beziehung zu den Aufgaben der allgemeinen Wissenschaft vom Leben. Forschungsberichte
aus der biologischen Station Plön 12:1–39
Zacharias O (1907) Das Süsswasserplankton. Teubner, Leipzig
Zacharias O (1909) Das Plankton als Lebensgemeinschaft. Unsere Welt 1:5–14
Part IV
Main Phases of the History
of the Concept “Ecology”
145
Literal Translation in German: Ökologie; in French: Écologie
The term “Oecologie” was coined by the German zoologist Ernst Haeckel in
1866 in his book Generelle Morphologie der Organismen.1 It derives from the
Greek “oikos” (oikos; house, household, also dwelling place, family) and “logos”
(logos; word, language, language of reason). “Oecologie”, later appearing as
“Ökologie” (from around 1890),2 was used to refer to the “whole science of the
relations of the organism to its surrounding outside world”3 or, to put it differently,
the science of the household of nature or the economy of organisms.4 The term was
taken up rather rapidly by some authors (e.g. by Semper 1868, p. 229), but more
than 20 years passed until it became widely used. It was not until 1885 that it was
A. Schwarz (*)
Institute of Philosophy, Technische Universität Darmstadt, Schloss, 64283 Darmstadt, Germany
e-mail: schwarz@phil.tu-darmstadt.de
K. Jax
Department of Conservation Biology, Helmholtz Centre for Environmental Research (UFZ),
Permoserstr. 15, 04318 Leipzig, Germany
e-mail: kurt.jax@ufz.de
Chapter 9
Etymology and Original Sources
of the Term “Ecology”
Astrid Schwarz and Kurt Jax
1 Haeckel 1866, Vol.1, p. 8 (footnote), 237, 238 (table); Vol. 2, p. 286. There is a persisting tale that
Thoreau used the word prior to this (see, for instance, Michel Serres in Revisiting the natural contract
2006). According to Walter Harding, editor of “The correspondence of Henry David Thoreau”, this
interpretation is a misreading that confused geology with ecology: “I must assume that, since geology
makes as much sense in the context as ecology does, geology must have been the word that Thoreau
intended”. (Harding 1965, p. 707 (emphasis in the original); see also Egerton 1977 or Acot 1982).
2 In the 1896 German version of his book on Ecological Plant Geography, Warming wrote
Ökologie while Dahl (1898), for example, still used the spelling Oekologie.
3 “Gesammte Wissenschaft von den Beziehungen des Organismus zur umgebenden Aussenwelt”,
Haeckel 1866 (p. 286, emphasis in original).
4 Haeckel 1870 (p. 365) originally referred only to animals: “[der] Haushalt der thierischen
Organismen” (“the household of animals”). This paper was a written version of his inaugural
lecture as a professor in Jena.
A. Schwarz and K. Jax (eds.), Ecology Revisited: Reflecting on Concepts, Advancing Science,
DOI 10.1007/978-90-481-9744-6_9, © Springer Science+Business Media B.V. 2011
146
A. Schwarz and K. Jax
used for the first time in a book title, namely in Reiter’s Die Consolidation der
Physiognomik als Versuch einer Oekologie der Gewaechse.
In English the term “Oecologie” was initially translated as “œcology”. Its first
use occurred, it seems, in the English translation of Haeckel’s book Natürliche
Schöpfungsgeschichte (The History of Creation5) in 1876 (cf. Bather 1902,6
p. 748; Benson 2000, p. 60). In the early 1890s, not least following a recommenda-
tion of the Madison Botanical Congress in 1893 (Madison Botanical Congress
1894, pp. 35–38), the double-letter was dropped in favour of the final English form
“ecology”. From that time on, the term “ecological” was widely used in publica-
tions7 and from 1904 the Botanical Gazette, a University of Chicago publication,
produced a column entitled Ecological notes.
However, the old wording still persisted for some years in the writings of a number
of authors and even more in dictionaries (Bessey et al. 1902; Bather 1902). For
example, the English translation of Warming’s (1895) seminal book Plantesamfund
was published in 1909 as Oecology of plants.
In French the word was also introduced as “oecologie” via the translation of
Haeckels “Natürliche Schöpfungsgeschichte” (Histoire de la création des êtres
organisés) in 1874. Its first use by French scientists (now as “écologie”) can be
traced definitively to the year 1900 in the context of plant ecology. Charles Flahault,
a botanist loosely associated with the Montpellier school, used the word in its
adjectival form in his “projet de nomenclature phytogéographique”.8 However,
despite being introduced by a famous zoologist and Darwinian apologist, the word
was not very successful in the French community of biologists; this might be due
to the fact that resistance to Darwinian ideas was much greater in France than in the
English-speaking world.9
In Russian the term “ekologia” was first introduced through an abridged translation
of Haeckel’s Generelle Morphologie der Organismen in 1869. In similar fashion to
its impact on the French and the English-speaking world, it was not influential in
the sense of leading to the founding of an ecological research programme or institu-
tions. In the late 1890s, this situation changed with the translation of Eugenius
Warming’s Plantesamfund into Oikologicheskaya Geografia Rastenii (1901) which
was highly influential, together with the translation of Grisebach’s work (1874,
1877): researchers now began to focus on groups of organisms and to develop a
synecological approach.
5 Haeckel 1876, Vol. II, p. 354.
6 Bather erroneously assumes that Haeckel also coined the word in the Natürliche
Schöpfungsgeschichte. Haeckel’s Generelle Morphologie der Organismen (1866), where the word
was in fact used first, was never published in English translation.
7 For instance A. S. Hitchcock, Ecological Plant Geography of Kansas (1898) or H. C. Cowles,
The ecological relations of the vegetation on the sand dunes of Lake Michigan (1899).
8 C. Flahault, Projet de nomenclature phytogéographique, Actes du 1er Congrès international de
botanique tenu à Paris à l’occasion de l’exposition universelle de 1900, Lons-le-Saunier, 1900,
p. 440, 445 (cf. Matagne 1999, p. 107).
9 Matagne 1999, p. 109; also Acot 1982, p. 106 ff.
147
9 Etymology and Original Sources of the Term “Ecology”
Brief Overview of Part IV to VII
The term “Oecologie” was coined in the second half of the nineteenth century.
However, Antecedents of an ecological idea in the sense of a modern science
and in contrast to natural history existed before it was described as “ecology”. Such
ideas were present, for example, in Alexander von Humboldt’s “physiognomic
system of plant forms”, Alfred R. Wallace’s “geography” of animal species, and
Charles Darwin’s “entangled bank”; they also existed in Louis Agassiz’ studies on
lakes and oceans. In 1866 “Oekologie” was introduced by the German zoologist
Ernst Haeckel. From the beginning he referred to the meaning of the Greek “oikos”
as “household”, suggesting that “ecology” is the science of the household of organ-
isms, i.e. of their relation to their biotic and abiotic surroundings. None of the defi-
nitions he offered denoted an existing research programme, nor was Haeckel’s aim
to develop such a programme. The term primarily filled an empty place within his
disciplinary system of zoology: that of “external physiology”. This search for an
order of the study of living beings was in line with contemporary efforts to find a
general system of biology. However, the new name “Oekologie” gave way to a
focus on the field as a self-conscious enterprise, despite varying local conditions
of early ecology according to nation-state and language area.
At the time a number of competing terms existed. “Ecology” was used in a
wide range of ways to refer to different domains of objects and phenomena. During
the late nineteenth and early twentieth century, what is nowadays called “ecology”
was also described as “ethology” or “biology” in a narrower sense. Likewise, different
approaches were taken towards delimiting the term “ecological” to terms such as
“physiological” and/or “sociological”.
In the first half of the twentieth century a stabilising of the concept occurred.
At the same time the formation of scientific societies, academic institutions and
publishing bodies accelerated, and the later subdivisions of ecology appeared on
the horizon. With the rise of systems theory in the 1940s and the arrival of the
environmental movement in the 1960s the concept broadened and ecology came
to be described as a “super-science”. “Ecology” in this sense served to blur the bound-
aries between scientific, philosophical and political knowledge, and at the method-
ological level there was a merging of facts and values, the epistemic and the social.
The border zones between scientific ecology and other fields became the
subject of highly controversial discussions. From this time on, “ecology” can refer to
a variety of completely disparate ideological doctrines and political stances.
The struggle to define ecology and its sub-disciplines has thus always been both
one of structuring and assessing the complex subject matter of “the interdepen-
dence of living and non-living nature” and a debate about the delimitation of insti-
tutional and social groups – in academia and beyond. This process is still ongoing
and will be of central importance in locating Ecology in the context of 21ST
century environmental sciences. It also has importance with respect to the
question of whether the domain of ecology is determined by method, object
or institution.
149
The neologism “Oecologie” was coined by German zoologist Ernst Haeckel in
1866. His intention in doing so, however, was not to establish a discipline of
“Oecologie” along with its own concepts, theories and practices. Haeckel himself
never engaged in “ecological” research, but rather invented the word to identify a
hitherto unnamed branch in his system of zoology.1 It was not until around the
1890s that ecology became a “self-conscious”2 enterprise. Prior to that, the term
served more as a focal point to denote certain activities that had been undertaken in
disciplines such as zoology, botany, physiology, geography and oceanography,
which in turn constituted the diverse roots of what would later be known as
“ecology”.
Haeckel presented the term “Oekologie” in several publications, providing a
variety of definitions for it.3 The elements contained in these definitions, along with
his characterization of the place of ecology within biology, are still in evidence in
debates about ecology today.
In Haeckels’s system of zoology, ecology refers to the external physiology of
organisms:
Die Physiologie theilen wir ebenfalls in zwei Disciplinen: I. Die Physiologie der
Conservation oder Selbsterhaltung (a. Ernährung, b. Fortpflanzung), II. die Physiologie der
Relationen oder Beziehungen (a. Physiologie der Beziehungen der einzelnen Theile des
Chapter 10
The Early Period of Word and Concept
Formation
Kurt Jax and Astrid Schwarz
1 This is highlighted by the fact that Haeckel first used the term “Oecologie” in a diagrammatic
representation before setting out to explicate it in words (Haeckel 1866, vol.1, p. 238).
2 Allee et al. 1949, pp. 19, 42.
3 Haeckel 1866, 1868, 1870.
K. Jax
Department of Conservation Biology, Helmholtz Centre for Environmental Research (UFZ),
Permoserstr. 15, 04318 Leipzig, Germany
e-mail: kurt.jax@ufz.de
A. Schwarz (*)
Institute of Philosophy, Technische Universität Darmstadt, Schloss, 64283 Darmstadt, Germany
e-mail: schwarz@phil.tu-darmstadt.de
A. Schwarz and K. Jax (eds.), Ecology Revisited: Reflecting on Concepts, Advancing Science,
DOI 10.1007/978-90-481-9744-6_10, © Springer Science+Business Media B.V. 2011
150
K. Jax and A. Schwarz
Organismus zu einander (beim Thiere Physiologie der Nerven und Muskeln); b. Oecologie
und Geographie des Organismus oder Physiologie der Beziehungen zur Aussenwelt).4
One perspective, then, places the focus on the interrelations between an organism
and its physical surroundings, including interactions with other organisms:
Unter Oecologie verstehen wir die gesammte Wissenschaft von den Beziehungen des
Organismus zur umgebenden Aussenwelt, wohin wir im weiteren Sinne alle ‚Existenz-
Bedingungen‘ rechnen können. Diese sind theils organischer, theils anorganischer Natur.5
An alternative perspective focuses on ecology as an “economy of nature”, implying
a reference to liberal economic theories as supposedly advocated – at least in
Haeckel’s view – by Darwin:
Die Oecologie der Organismen, die Wissenschaft von den gesammten Beziehungen der
Organismen zur umgebenden Außenwelt, zu den organischen und anorganischen
Existenzbedingungen; die sogenannte ‚Oekonomie der Natur’, die Wechselbeziehungen
aller Organismen, welche an einem und demselben Orte mit einander leben, ihre Anpassung
an die Umgebung, ihre Umbildung durch den Kampf um’s Dasein.6
Unter Oecologie verstehen wir die Lehre von der Oeconomie, von dem Haushalt der
thierischen Organismen. Diese hat die gesammten Beziehungen des Thieres sowohl zu
seiner anorganischen, als zu seiner organischen Umgebung zu untersuchen, vor allem die
freundlichen und feindlichen Beziehungen zu denjenigen Pflanzen und Thieren, mit denen
es in directe oder indirecte Berührung kommt; oder mit einem Worte alle diejenigen ver-
wickelten Wechselbeziehungen, welche Darwin als die Bedingungen des Kampfes um´s
Dasein bezeichnet.7
Haeckel’s definitions explicitly excluded the spatial (topographical and geo-
graphical) dimension, which he reserved for what he called “chorology” (Arealkunde,
4 “We divide physiology also into two disciplines: I. the physiology of conservation or self-
preservation (a. nutrition, b. reproduction), II. the physiology of relations (a. physiology of the
relations of parts of the organism to each other (meaning, for animals, the physiology of nerves
and muscles); b. ecology and geography of the organism or physiology of the relations with the
external world)” (Haeckel 1866, vol. 1, p. 237).
5 “By ecology we mean the whole science of the relations of the organism to its surrounding out-
side world, which we may consider in a broader sense to mean all ‘conditions of existence’. These
are partly of an organic nature and partly of an inorganic nature” (Haeckel 1866, vol. 2, p. 286
(emphasis in original)).
6 “The ecology of the organisms, the science of the whole relations of organisms to their surrounding
world, towards the organic and inorganic conditions of existence; the so-called ‘economy of
nature’, the interrelations of all organisms which live in one and the same place, their adaptations
to their environment, their transformation through the struggle for existence” (Haeckel 1868,
p. 539 (emphasis in original)).
7 “By ecology, we mean the science of the economy, of the household of animal organisms. This
has to study the entirety of relations of the animal both to its inorganic and its organic environ-
ment, in particular the benign and hostile relations with those plants and animals with which it
comes directly into contact; or, to be concise, all those intricate interrelations which Darwin calls
the struggle for existence” (Haeckel 1870, p. 365).
15110 The Early Period of Word and Concept Formation
biogeography).8 This spatial dimension, however, became one of the basic pillars
of the concept of ecology.
The positioning of ecology as part of physiology was part of Haeckel’s search
for a structure of biology that was logically related to Darwin’s theory of evolution,
from which he also drew insights about the importance of those relations of organisms
which he called “ecological”.9
Figure 10.1a shows the scheme of the main branches of zoology, in which Haeckel
placed ecology and chorology in 1866: ecology and chorology are situated within the
“physiology of the relations of organisms to their outside world”. They are part of the
“Relations-Physiologie der Thiere” (physiology of relations of animals), the other
part of which is devoted to the relations of the parts of the (animal) body to each other.
This physiology of relations was set in contrast to the “Conservations-Physiologie”
(physiology of conservation, meaning the physiology by which the organism keeps
itself alive and reproduces). As already noted critically by Tschulok (1910, pp. 141
ff) the specific placement of ecology and, especially, chorology (geography and
topography as a part of physiology) does not follow any clear logic or explicit criteria.
It is not obvious, for example, why the physiology of muscles and nerves (which he
provides as examples of the “relations of the individual parts of the animal body to
each other”) is closer to the physiology of the organism’s relations to its surroundings
than they are to the physiology of internal metabolism or reproduction. It seems that
Haeckel‘s desire to find a consistent scheme of clear-cut dichotomies into which the
different parts of biology (or here, specifically zoology) could be ordered, was a
dominant force in the design of his scheme. This is evident also from the ease with
which Haeckel rearranged parts of the scheme. Figure 10.1b shows a second visual-
ization of the divisions of zoology, published in 1902.10 Here, parts of what had previ-
ously appeared as “relations physiology” (Relations-Physiologie), in particular the
“relations of the individual parts of the animal body to each other”, have been shifted
to the other part of physiology, the physiology of working functions (Physiologie der
Arbeitsleistungen), without any clear indication as to why – or indeed to which specific
category – they were moved. While keeping the overall number of branches of zool-
ogy constant (eight at the lowest level), chorology and ecology now become more
8 “Unter Chorologie verstehen wir die gesammte Wissenschaft von der räumlichen Verbreitung der
Organismen, von ihrer geographischen und topographischen Ausdehnung über die Erdoberfläche”
(Haeckel 1866, Vol. 2, p. 287, emphasis in original). (By chorology we mean the whole science of
the spatial distributions of the organisms, of their geographical and topographical extension over
the surface of the earth).
9 See Stauffer (1957). During the nineteenth and the first half of the twentieth century several
publications attempted to link the different branches of biology in a consistent scheme; in addition
to that of Haeckel (1866), these included in particular Burdon-Sanderson (1893), Wasmann
(1901); Tschulok (1910), Gams (1918), and Du Rietz (1921).
10 Although the figure is taken from an 1870 “reprint” of Haeckel’s paper in a collection of his popu-
lar writings, it was not included in the original paper. Also, the terminology and the ordering used
there (1870) is intermediate between those of the two schemes displayed in (Fig. 10.1).
152
separate as the only parts of the physiology of relations (with chorology now also
including the subject species’ migrations). Obviously, in both representations the field
“Oecologie” followed (at most) the internal logic of Haeckel’s system and was never
intended as a vehicle to develop a consistent concept of ecology in the sense of a new
research programme.
Nevertheless, the concept of ecology as the “external physiology” of organisms
was taken up early on,11 and by the turn of the twentieth century at the latest, the
11 This is the case especially in the German-speaking world, where the reference to the physiological
tradition was more important than in the Anglo-Saxon world (e.g. Trepl 1987, p. 26). One of the
first authors to develop a research programme in this sense was Karl August Möbius, with his case
study of the economy of oyster beds (Möbius 1877).
Fig. 10.1 Ernst Haeckel’s ideas about the subdivisions of zoology. (a) From Haeckel (1866)
(Vol. 1, p. 238), (b) from Haeckel (1902), p. 29. See text for explanation
K. Jax and A. Schwarz
15310 The Early Period of Word and Concept Formation
early protagonists of ecology such as Carl Semper, A.F. Wilhelm Schimper and
Eugenius Warming, as well as Frederic Clements and Henry Chandler Cowles,
succeeded in developing it into a research programme. The emphasis on the biotic
interrelations of organisms as well as a consideration of the whole “economy
of nature” were part of these and other research programmes. Stabilizing the range
of meanings of the concept of ecology and its varied transformations was one of the
key concerns of ecologists in the early stages of the discipline’s formation.12
References
Allee WC, Emerson AE, Park O, Park T, Schmidt KP (1949) Principles of animal ecology.
Saunders, Philadelphia
Burdon-Sanderson JS (1893) Inaugural address. Nature 48:464–472
Du Rietz GE (1921) Zur methodologischen Grundlage der modernen Pflanzensoziologie. Adolf
Holzhausen, Wien
Gams H (1918) Prinzipienfragen der Vegetationsforschung. Ein Beitrag zur Begriffsklärung und
Methodik der Biocoenologie. Vierteljahresschr. Naturf Gesellsch Zürich 63:293–493
Haeckel E (1866) Generelle Morphologie der Organismen. Georg Reimer, Berlin
Haeckel E (1868) Natürliche Schöpfungsgeschichte: gemeinverständliche wissenschaftliche
Vorträge über die Entwickelungslehre im Allgemeinen und diejenige von Darwin, Goethe und
Lamarck im Besonderen, über die Anwendung derselben auf den Ursprung des Menschen und
andere damit zusammenhängende Grundfragen der Naturwissenschaft. Reimer, Berlin
Haeckel E (1870) Über Entwicklungsgang und Aufgabe der Zoologie. Jenaische Z Med Naturwiss
5:353–370
Haeckel E (1902) Gemeinverständliche Vorträge und Abhandlungen aus dem Gebiete der
Entwickelungslehre. Emil Strauß, Bonn
Möbius KA (1877) Die Auster und die Austernwirtschaft. Wiegandt, Hempel & Parey, Berlin
Stauffer RC (1957) Haeckel, Darwin, and ecology. Q Rev Biol 32:138–144
Tschulok S (1910) Das System der Biologie in Forschung und Lehre. Eine historisch-kritische
Studie. Gustav Fischer, Jena
Wasmann E SJ (1901) Biologie oder Ethologie? Biols Zentralbl 21:391–400
12 See Jax, Chap. 12, this volume.
155
Chapter 11
Competing Terms
Kurt Jax and Astrid Schwarz
Up until the early twentieth century, competing terms for “ecology” were “natural
history”, “biology”, “bionomics”, and “ethology”.
Natural history is one of the roots of ecology and played an important role in the
emergence of ecology. Haeckel noted in 1870 (p. 365): “Ecology (often also inap-
propriately called biology in the narrower sense) has, up to now, constituted the
main component of so-called ‘natural history’ in the usual sense of this word.”1
Although he places ecology (as “external physiology”) in the context of a mod-
ern system of biology,2 he nevertheless acknowledges the importance of natural
history as a fundamental root of ecology. The traces of a natural history in the sense
of “structuring visible nature”3 never disappeared entirely in ecology. This is
despite the fact that, by the turn of the twentieth century, natural history was largely
identified with a rather indiscriminate and “unscientific” way of collecting data
about natural phenomena. However, this diagnosis was already a result of the steady
boundary work going on in the emerging laboratory-based biological sciences. In
fact, the natural history of the nineteenth century never was merely a matter of
haphazard sampling and describing; rather, it had a very specific methodology and
a very specific set of theoretical questions.4 Such questions included the issue of
1 “Oecologie (oft unpassend auch als Biologie im engeren Sinne bezeichnet) bildete bisher den
Hauptbestandtheil der sogenannten ‚Naturgeschichte’ in dem gewöhnlichen Sinne des Wortes.”
2 In his writings Haeckel always refers specifically to zoology.
3 Translated from Foucault 1974, p. 177.
4A detailed discussion of the changing rationality of natural history through the centuries is given in
Cultures of Natural History (1996), edited by Jardine, N., E.C. Spary and J.A. Secord; R. Kohler (2006)
presented a detailed study on specific scientific practices in natural history in the nineteenth century
through to the 1950s, while D. Takacs shows in The Idea of Biodiversity. Philosophies of Paradise (1996)
that natural history is very present in the modern biosciences.
K. Jax
Department of Conservation Biology, Helmholtz Centre for Environmental Research (UFZ),
Permoserstr. 15, 04318 Leipzig, Germany
e-mail: kurt.jax@ufz.de
A. Schwarz (*)
Institute of Philosophy, Technische Universität Darmstadt, Schloss, 64283 Darmstadt, Germany
e-mail: schwarz@phil.tu-darmstadt.de
A. Schwarz and K. Jax (eds.), Ecology Revisited: Reflecting on Concepts, Advancing Science,
DOI 10.1007/978-90-481-9744-6_11, © Springer Science+Business Media B.V. 2011
156
explaining the distribution and origin of natural variety and the mechanisms of
nature’s perceived order.5
The identification of ecology with “classical” natural history was strongly
rejected by most early ecologists as being too broad (e.g. Wheeler 1902). In
response, however, to what he perceived as an overemphasis of laboratory biology
at the cost of observing animals in the field, Charles Elton wrote of ecology in 1927
in terms of it being “scientific natural history” (Elton 1927, p. 1). Shelford (1937
p. 32, FN 1) also noted that “[t]he term is applied to those phases of natural history
and physiology which are organized into a science, but does not include all the
unorganizable data of natural history”.
In the second half of the nineteenth century and into the early twentieth century,
“biology” was used (especially in Germany) both in the broad sense common today
and in a narrower sense, designating a concept that was close to, if not identical
with, Haeckel’s definition of ecology.6 This dual use of the word and its application
to “ecology” (or sometimes also “ethology”) was criticised for several reasons, not
least because it was seen as leading to confusion (Haeckel 1866, 1870; Wasmann
1901;7 Dahl 18988). Wasmann, who on the one hand criticised this dual use, tried
on the other to save and refine the notion of “biology” in its narrower sense. For
him, it included “the external activities of life which pertain to organisms as indi-
viduals and at the same time regulate their relations to other organisms and to the
inorganic conditions of existence”.9 It thus comprised the external habits of organ-
isms (such as feeding and reproduction), their interactions, and their conditions of
existence. In his view, this concept was broader than that of “ethology” and even
more so than that of Haeckel’s “ecology”.
5 The questions posed here were also “why” and not only “how”. The methodology of natural history,
as the term denotes, was a heavily historical one, explaining the specific patterns and processes
found in nature in a nevertheless systematic way (Trepl 1987, p. 46). Farber (1982, p. 150 f.) sees
natural history and “scientific” physiology as parallel traditions in the nineteenth century, with
natural history guided by theoretical questions (e.g. the relations between classification, morphol-
ogy and history) and culminating, by the middle of the century, in evolutionary theory.
6 Haeckel saw ecology as a substitute for what he perceived as the meaning of “biology in the
narrow sense”.
7 In spite of the criticism that he himself also raised with regard to this issue, Wasmann argued in favour
of retaining the term “biology” in the narrower sense – rejecting both “ethology” and “ecology”.
8 Dahl (1898, p. 121 f.) argued: “Hat man doch bisher nicht einmal einen Namen fur dieses Gebiet
gefunden, der allgemein anerkannt wurde. Man nannte es fruher Biologie. Nachdem aber diese
Bezeichnung im weitesten Sinne auf die Erforschung aller Lebewesen in Anwendung gekommen ist
und die Zellforschung im Speziellen sich Biologie nennt, müssen wir als die minder Bekannten und
Geachteten das Feld räumen”. (“Thus far, a name that might meet with general agreement has not even
been found for the field. It used to be called biology. But since this name has now been applied in the
broadest sense to the investigation of all living beings and since cytological research in particular calls
itself biology, we, as those who are less well-known and respected, are compelled to beat a retreat”).
9 “…[die] äußeren Lebensthätigkeiten, die den Organismen als Individuen zukommen, und die zugleich
auch ihr Verhältnis zu den übrigen Organismen und zu den anorganischen Existenzbedingungen regeln”
(Wasmann 1901, p. 397).
K. Jax and A. Schwarz
157
The use of “biology” rather than “ecology” persisted in the writings of the community, a
fact that was noted and criticised by, among others, Sinai Tschulok in 1910 (p. 211 f.):
“Es ist sehr zu bedauern, daß selbst in wissenschaftlichen Werken die Ökologie
noch sehr häufig als Biologie bezeichnet wird. Denn einmal ist Biologie die
Gesamtwissenschaft von den Lebewesen, ein anderes Mal ein Teil davon, d.h.
eine bestimmte Betrachtungsweise […]. Noch schlimmer ist es, wenn unter
Biologie etwas mehr als Ökologie verstanden werden will und damit doch nicht die
gesamte Lehre von den Organismen gemeint ist”.10
In the second half of the nineteenth century another, narrower meaning of biology
in the sense of “cytology” (later relating also to “microscopic knowledge of organ-
isms”) had been established, especially in France and Belgium (Dahl 1898;
Wasmann 1901; Wheeler 1926). This in turn led to the necessity to coin a new term
for the “science of the ways of living of animals and plants” (Wasmann 1901, p. 394),
namely, “ethology”.11 Ethology was a term preferred especially by zoologists
(e.g. Dahl 1898; Wheeler 1902). Dahl saw ecology as part of ethology, together
with what he called “trophology” (pertaining to the food of animals). Both Dahl
and Wheeler saw “ecology” as too narrow, arguing that it referred (at least through
the direct connotation of the word “oikos”) mainly to “dwelling” (“Aufenthalt”,
Dahl 1898, p. 122) or “habitat” (Wheeler 1902, p. 973). Wheeler also opted for the
term “ethology” to describe the whole complexity of animal life, including (in
contrast to plant life) social and psychological phenomena of the organisms’ rela-
tions, while this emphasis was not a major interest of Dahl’s. Like Dahl, Wheeler
too saw ecology as a part of ethology. Attempts to substitute “ecology” for “ethology”,
however, were not a great success. By the 1920s, if not earlier, the word “ecology”
had already become the dominant term within zoology as well, while ethology had
soon become restricted to the behaviour of animals in terms of their individual
social interactions. Carpenter’s “Ecological Glossary” from 1938 doesn’t even
mention “ethology”. However, in the same period the German biologist Jakob von
Uexküll propagated a “theory of environment” (Umweltlehre), managing to ignore
both the labels “ecology” and “ethology” in the process.12
10 “It is highly regrettable that even in scientific works ecology is still very frequently referred to
as biology. For biology is at once the overall science of living being as well as a part of it, i.e. a
particular perspective... The matter becomes even worse when biology is intended to denote some-
thing more than ecology and thus not the general study of organisms”.
11 The first author to use “ethology” in this sense was Isidore de Geoffrey St. Hilaire (1854)
(Wheeler 1926; van der Klauuw 1936a; Jahn and Sucker 2000). See van der Klaauw 1936a, p. 140
for the precise wording of this first description of “ethology”.
12 “Die Umweltlehre besteht (…) aus zwei Hauptpunkten. Neben der Anerkennung der planer-
zeugten Umwelten fordert sie die Anerkennung des Zusammenhanges aller Umwelten in einer
allumfassenden Planmäßigkeit.” (“The theory of the environment consists of (...) two main points.
In addition to acknowledging plan-based environments it demands acknowledgement of the inter-
relatedness of all environments in an all-encompassing orderliness”. Uexküll 1929, p. 45).
Uexküll’s specific concept of Umwelt (not really captured in today’s colloquial term “environ-
ment”) was at once nomothetic and idiographic, in the sense of depending on the specific
individual (biological subject).
11 Competing Terms
158
“Bionomics”, was another, less commonly used concept that “competed” with
ecology,13 and still appears to be the least defined of these concepts even today.
In 1910 Dahl – in contrast to his earlier writings – saw ethology as being on an equal
level with ecology which, together with psychology, constitutes animal bionomics,
while he considered plant bionomics as being identical with plant ecology (Dahl
1910, p. 3). For Friederichs (1930) bionomics was the science of the specific “laws
of life” (Lebensgesetzlichkeit; p. 11) of a species as these are manifested externally,
but was still closely related to ecology. Carpenter (1938) defines bionomics as “aute-
cology (used more loosely)” (p. 44); for Schaefer (2003) it is “the science of the way
of living of a species, also called ‘biology in the narrower sense’14; and for Lincoln
et al. (1998, p. 41) it is a synonym of ecology: “Ecology; the study of organisms in
relation to their environment”. Thus, the naming competition seems to be ongoing
even now, despite the fact that ecology nowadays is well institutionalised.
References
Carpenter RJ (1938) An ecological glossary. Kegan Paul, Trench, Trubner & Co., London
Dahl F (1898) Experimentell-statistische Ethologie. Verhandlungen der Deutschen Zoologischen
Gesellschaft, Jahresversammlung 1898 in Heidelberg:121–131
Dahl F (1910) Anleitung zu zoologischen Beobachtungen. Quelle & Meyer, Leipzig
Elton C (1927) Animal ecology. Sidgwick & Jackson, London
Farber PA (1982) The transformation of natural history in the nineteenth century. J. Hist. Biol.
15:145–152
Foucault M (1974) Die Ordnung der Dinge. Suhrkamp, Frankfurt/Main
Friederichs K (1930) Die Grundfragen und Gesetzmäßigkeiten der land- und forstwirtschaftlichen
Zoologie, insbesondere der Entomologie. Erster Band: Ökologischer Teil. Paul Parey, Berlin
Haeckel E (1866) Generelle Morphologie der Organismen. Georg Reimer, Berlin
Haeckel E (1870) Über Entwicklungsgang und Aufgabe der Zoologie. Jenaische Zeitschrift für
Medizin und Naturwissenschaften 5:353–370
Jahn I, Sucker U (2000) Die Herausbildung der Verhaltensbiologie. In: Jahn I (ed) Geschichte der
Biologie, 3 edn. Spektrum, Akademischer Verlag, Jena, pp 580–600
Jardine N, Secord JA, Spary EC (eds) (1996) Cultures of natural history. Cambridge University
Press, Cambridge
Klaauw C, J. van der (1936) Zur Geschichte der Definitionen der Ökologie, insbesonders aufgrund
der Systeme der zoologischen Disziplinen. Sudhoffs Archiv für Geschichte der Medzin und
der Naturwissenschaften 29:136–177
Kohler RE (2006) All Creatures: Naturalists, Collectors and Biodiversity 1850-1950 Princeton
University Press, Princeton, New Jersey
Lincoln R, Boxshall G, Clark P (1998) A dictionary of ecology, evolution and systematics.
Cambridge University Press, Cambridge
13 Wheeler (1926) mentions E. Ray Lankester as one who used the term in 1889, while Jahn &
Sucker (2000) refer to J. Wilhelm Haake as the first to use it, Lankester and Haake being students
of Haeckel. “Zoonomie”, describing the (natural) history of animals, can already be found in the
early nineteenth century (see van der Klauuw 1936a, p. 137 f.).
14 “Lehre von der Lebensweise einer Art, wird auch als ’Biologie im engeren Sinne’ bezeichnet”
(Schaefer 2003, p. 50).
K. Jax and A. Schwarz
159
Schaefer M (2003) Wörterbuch der Ökologie, 4 edn. Spektrum Akademischer Verlag,
Heidelberg
Shelford VE (1937) Animal communities in temperate America, 2 edn. University of Chicago
Press, Chicago
Takacs D (1996) The idea of biodiversity: philosophies of paradise. John Hopkins University
Press, Baltimore & London
Trepl L (1987) Geschichte der Ökologie. Vom 17. Jahrhundert bis zur Gegenwart. Athenäum,
Frankfurt/Main
Tschulok S (1910) Das System der Biologie in Forschung und Lehre. Eine historisch-kritische
Studie. Gustav Fischer, Jena
Uexküll Jv (1929) Welt und Umwelt. Aus deutscher Geistesarbeit 5:20–26, 36–46
Wasmann ESJ (1901) Biologie oder Ethologie? Biologisches Zentralblatt 21:391–400
Wheeler WM (1902) ‘Natural history’, ‘oecology’ or ‘ethology’? Science 15:971–976
Wheeler WM (1926) A new word for an old thing. Q. Rev. Biol. 1:439–443
11 Competing Terms
161
Chapter 12
Stabilizing a Concept
Kurt Jax
The concept of ecology emerged well before Haeckel’s coining of the word in the
nineteenth century, when the logical possibility – and with it the quest – arose to
understand the patterns of plant and animal distribution as a consequence of their
immediate environmental relations. However, the formation of ecology into a self-
conscious discipline1 – that is, a systematic enterprise engaged in by researchers
who described themselves as doing “ecological” work – postdated Haeckel’s defi-
nition of ecology by some two decades. In between lay the struggle to mark the
boundaries of the concept of ecology, and this phase of stabilizing ecology lasted
well into the first decade of the twentieth century.
Henry Chandler Cowles, for example, was asked to report to the American
Association for the Advancement of Science (AAAS) about “[t]he work of the year
1903 in ecology”. He began his report by stating: “It is more than impossible to do
such a task for ecology, since the field of ecology is chaos. Ecologists are not agreed
even as to fundamental principles or motives; indeed, no one at this time, least of all
the present speaker, is prepared to define or delimit ecology”. Cowles (1904, p. 879).
Ecology at the turn of the twentieth century was, in fact, still struggling to find its place
within the field of tension between its basic roots and constituent elements, namely
natural history and biogeography on the one hand, and modern biology (specifically,
physiology) on the other – to name only the most prominent ones.2
1 An expression used first by Allee et al. 1949 and then by McIntosh 1985.
2 Oceanography and limnology also contributed to the formation of ecology, albeit often through
a somewhat separate discourse. See, e.g. Schwarz 2003.
K. Jax ()
Department of Conservation Biology, Helmholtz Centre for Environmental Research (UFZ),
Permoserstr. 15, 04318 Leipzig, Germany
e-mail: kurt.jax@ufz.de
A. Schwarz and K. Jax (eds.), Ecology Revisited: Reflecting on Concepts, Advancing Science,
DOI 10.1007/978-90-481-9744-6_12, © Springer Science+Business Media B.V. 2011
162
K. Jax
The tension between the more descriptive, heavily context-dependent approaches
of natural history and those of physiology as a science based largely on the (controlled)
experiment has remained in evidence throughout the entire history of ecology. This
tension proved (and still proves) to be both productive and problematic. It trans-
formed ecology into an approach which not only united different ways of looking at
living nature but also developed powerful centrifugal forces, resulting in attempts to
“purify” the new discipline and rid it of particular strands of its tradition, particular
that of natural history.3
The strong emphasis placed on physiology by many early ecologists was also a
means to professionalize a certain part of biology, one which lay beyond the main-
stream of laboratory biology and was predominant at the end of the nineteenth and the
beginning of the twentieth century. The upcoming field of ecology at that time was
frequently seen as a passing fad by many scientists. Bringing physiology from the lab
to the field, not only made use of the established reputation of exact and precise bio-
logical methodology but also served to separate the “mere naturalists” from the “pro-
fessional ecologists” (Hagen 1986; Cittadino 1980). Major proponents of the young
science (e.g. Carl Semper, Andreas Schimper, Henry C. Cowles, Frederic Clements,
Victor Shelford, Richard Hesse) subscribed to this view.4
The Madison Botanical Congress, struggling with the proper definition (and
orthography) of “ecology”, dealt with this issue in the context of “plant physiology” –
or, as it was sometimes also named at that time, “vegetable physiology”.5 Frederic
Clements (1905, p. 2) stated: “There can be little question in regard to the essential
identity of physiology and ecology. (…) Ecology has been largely the descriptive
study of vegetation; physiology has concerned itself with function”, urging a
merger of the two, which “will combine the good in each, and at the same time
eliminate superficial and extreme tendencies” (ibid. p. 3).
Many early animal ecologists also perceived ecology in this way. For the
German biologist Carl Semper, ecology was part of a research programme to validate
Darwin’s evolutionary theory by investigating “the influence of temperature, light,
heat, humidity, nutrition, etc. on the living animal” (Semper 1868, p. 228f.).6 Victor
Shelford, quoting approvingly from Semper’s book on animal ecology (Semper
1880 – the first book on animal ecology)7 wrote in 1913: “At present ecology is that
part of general physiology which deals with the organism as whole, with its general
life processes, as distinguished from the more specialized physiology of organs, and
which also considers the organism with particular reference to its usual environment”.8
3 See Simberloff (1980) for a discussion of these endeavours.
4 See also Kohler (2002) for a historical account of the shifting lab-field boundary in biology.
5 Madison Botanical Congress (1894, pp. 35ff).
6 “... die Einflüsse des Lichtes und der Wärme, des Feuchtigkeitsgrades, der Nahrung etc. auf die
lebenden Thiere”. See also Chap. 19; for a discussion of Semper and the physiological tradition
in early German ecology.
7 The English versions of Semper’s book – with slightly different titles in the UK and the USA –
were published in 1881.
8 Shelford (1913, p. 1 (emphasis in original)).
163
12 Stabilizing a Concept
And in 1927, Richard Hesse wrote: “[E]cology [is] merely a continuation of and a
supplement to physiological anatomy […]; the conditions of the environment are
incorporated into the theoretical linking of the individual processes”.9
The physiological tradition of ecology (and, in particular, of what was later
termed “autecology”) had already been prepared by a school of German plant
physiologists around Simon Schwendener, Gottlieb Haberlandt and Ernst Stahl,
who – using an evolutionary perspective – sought to link morphology with physiology
in “Physiologische Pflanzenanatomie” (physiological plant anatomy).10 This
approach investigated the morphological features of plants with respect to their
functional (adaptive) significance, bringing laboratory science to the field.11 An
important step, however, which had a decisive influence on the stabilization of the
concept of ecology, was the merging of the physiological perspective in ecology
with a biogeographical one. This entailed, at the same time, a shift from the envi-
ronmental relations (and adaptedness) of single species or two-species interactions
(such as parasitism and mutualism) to that of communities. It was a step first taken
systematically by Eugenius Warming in Denmark and Andreas Schimper in
Germany (Warming 1896; Schimper 1898). They completed the transformation of
Alexander von Humboldt’s physiognomic plant geography (1807) from a partly
aesthetic, partly scientific perspective to an exclusively scientific one, a transforma-
tion that had already been prefigured some decades earlier by the work of August
Grisebach (1838). With Warming and Schimper, the programme of making physiology
the basis of an ecological plant (and animal) geography (as opposed to a merely
taxonomic or historical one) thus became a major aim of early ecology. Warming
(1896) wrote: “A purely physiognomic system has no scientific significance; only
when physiognomy is explained physiologically and ecologically does it gain this
significance”.12 and a few pages prior to this: “Why do plants unite into communities
and why do they have the physiognomy they have?”.13
Warming, who can more plausibly be considered a founder of ecology than
Haeckel, wrote this in his well received book entitled Lehrbuch der ökologischen
Pflanzengeographie (Textbook of ecological plant geography). Here he distin-
guished “ökologische Pflanzengeographie” (“ecological plant geography”) from
“floristischer Pflanzengeographie” (“floristic plant geography”):
“Die ökologische Pflanzengeographie […] belehrt uns darüber, wie die Pflanze und die
Pflanzenvereine ihre Gestalt und ihre Haushaltung nach den auf sie einwirkenden Faktoren,
9 Hesse (1927, p. 944): “[...D]ie Ökologie [ist] nur eine Fortführung und Ergänzung der physiolo-
gischen Anatomie […]; es werden die Bedingungen der Umwelt mit einbezogen in die gedankliche
Verknüpfung der Einzelvorgänge.”
10 This was also the title of Haberlandt’s (1884) seminal book on the new approach.
11 See especially Cittadino (1990) for a description of these schools.
12 Warming (1896, p. 5): “Ein rein physiognomisches System hat keine wissenschaftliche Bedeutung:
erst wenn die Physiognomie physiologisch und ökologisch begründet wird, erhält sie eine solche”.
13 Ibid., p. 3 (emphasis in the original): “Weshalb schließen sich Arten zu bestimmten Gesellschaften
zusammen und weshalb haben diese die Physiognomie, die sie besitzen?”. For the transformation of
Humboldts’s plant physiognomy into ecology see (Trepl 1987, pp. 104ff; Schwarz 2003, pp. 28ff).
164
K. Jax
z.B. nach der ihnen zur Verfügung stehenden Menge von Wärme, Licht, Nahrung und
Wasser u.a. einrichten”.14
Extending the notion of “external physiology” from the physiology of organisms to
the physiology of communities proved to be a decisive step for ecology. It was this
programme that Schimper, Clements, Shelford and Hesse also pursued (Schimper
1898; Clements 1905; Shelford 1913; Hesse 1924).
In its stringent version, however, the programme of a “purely” physiological and
experimental ecology never fulfilled its promises, at least not within terrestrial ecology
and particularly with regard to its emphasis on the experimental method (Hagen
1988). Description, comparison and classification – the basic tools of natural history –
remained essential for ecological work and theory building. Indeed Warming’s
book itself (and many of Clements’ empirical writings) deals not only with abiotic
factors and experimental approaches to investigating how plants relate to these factors,
but also with interactions among organisms and the dynamics and classification of
whole plant communities.15 Warming integrated a physiognomic perspective
(derived from earlier plant geography), a physiological perspective concerned with
the “household” of plants (and plant societies) in relation to their abiotic environment,
and a strong emphasis on the interactions among organisms as a structuring element
of communities.
Thus around 1900 the main “ingredients” of the concept of ecology were all in
place – not just as mere definitions but as practice:16
The external physiology of organisms as an undisputed core element of ecology, •
involving
The interactions of organisms with abiotic factors as well as their interactions •
with each other
The set of relations between external physiology and the local and global (or at •
least regional) distribution of organisms
An extension of the basic objects of ecology from the individual organism to •
whole communities17
14 Ibid., p. 2, emphasis in original: “Ecological plant geography [...] teaches us how plants and
plant communities adjust their forms and their housekeeping to the factors that affect them,
e.g. the available amount of heat, light, nutriment, water, and so forth”.
15 The original Danish title (Warming 1895) was “Plantesamfund”, meaning “Plant society” or
“Plant community”; see Chap. 23 for a detailed account of Warming’s notion of ecology.
16 In terms of his pure verbal definition of ecology, Warming simply refers to Haeckel’s description
of ecology as the “Wissenschaft von den Beziehungen der Organismen zur Aussenwelt” (“the science
of the organisms’ relations to the exterior world”; Warming 1896, p. 2, footnote 1). However, as
described above, his concept of ecology is much more elaborate.
17 Different notions arose concerning the nature of “wholes” (or units) as objects of ecology, based
partly on physiognomic and “statistical” concepts and partly on more relational or even functional
ones (see Jax 2006 and also Chap. 4). Warming’s position, for example, came close to that formu-
lated later on by H.A. Gleason as the “individualistic concept of the plant community” (Gleason
1917, 1926). “Populations” and “ecosystems” were later arrivals as units of ecological investigation
(see also Chap. 14).
165
12 Stabilizing a Concept
Natural history remaining an indispensable part of ecology• 18
Despite this, the exact boundaries of the concept of “ecology” continued to be
contested and to “oscillate”19; to a lesser extent, this remains the case today. There
was considerable variation in the degree of emphasis placed on the many different
“ingredients” of the ecology concept. This became particularly apparent in the
many debates about the contexts in which the words “ecology” and “ecological” are
applied and in disagreements over how to demarcate ecology from other scientific
fields.20 A few important strands of these debates are outlined in the remainder of
this chapter.
British plant ecologist Arthur Tansley embraced a view of ecology similar to
Warming when he stated in his presidential address as the first president of the new
British Ecological Society in 1914: “We claim for ecology that it is before all things
a way of regarding the plant world, that it is par excellence the study of plants for
their own sakes as living beings in their natural surroundings, of their vital relations
to these surroundings and to one another, of their social life as well as their individual
life.” Tansley (1914, p. 195 (emphasis in original)). Others, however, like botanist
Paul Jaccard, attempted to introduce a distinction between ecology, chorology and
the sociology of plants and to restrict the meaning of “ecology”. He proposed to “...
use ecology only for the edaphic, physiographic, and climatic factors” (Jaccard in
Flahault and Schröter 1910, p. 21) and suggested using “chorology” for the
geographic relations of plants and “sociology” for the biotic interactions between
plants. His definition, entailing a narrower use of “ecology” than that proposed by
Haeckel and others, was not, however, widely accepted.
At the same time, the “plant sociology”, which Jaccard sought to restrict to
biotic relations alone, was often made synonymous with ecology – in particular
with synecology (as in Nichols 192321; Tansley 1920) or was conceived of even
more broadly, including chorology.22 Josias Braun-Blanquet wrote in the introduc-
tion to his textbook on plant sociology: “Plant sociology, the science of communities
or the knowledge of vegetation in the widest sense, includes all phenomena which
18 Albeit only its “organized parts”, as Victor Shelford emphasized, among others: “The term ecology
is applied to those phases of natural history and physiology which are organized into a science, but
does not include all the unorganizable data of natural history”. Shelford (1913, p. 32, footnote 1).
Similarly, more than a decade later, Charles Elton defines ecology as “scientific natural history”
(Elton 1927, p. 1).
19 Schwarz (2003, pp. 254ff), focusing mainly on aquatic ecology, describes ecology as oscillating
between three basic concepts, which she labels “microcosm”, “niche” and “energy”, representing
in turn a physiognomic, a physiological, and a functional approach.
20 See contributions in Part VII where the border zones between ecology and other scientific fields
are discussed in more detail.
21 “The study of plant communities in their relation to environment comprises the field of what
might be called ecological plant sociology; more commonly it has been called synecology”.
Nichols (1923, p. 11 (emphasis in original)).
22 Which is in fact also contained in Warming’s concept of ecology.
166
K. Jax
touch upon the life of plants in social units”.23 The “life of plants in social units”
meant for him: “the organization or structure of the community (...), synecology: the
study of the dependence of plant communities upon one another and upon the
environment, synergetics (development of communities) (...), synchorology (geographic
distribution of communities) (...), sociological classification (systematics) (...)”.24
Some later authors, however, used the word “plant sociology” (even more common:
“phytosociology”) for the classification of plant assemblages only (see Friederichs
1963). Thus “phytosociology” is defined in a British dictionary of ecology as: “The
classification of plant communities based on floristic rather than life-form or other
considerations”.25
“Sociological” is used in a much more restricted way in animal ecology, where
it is usually (though not always) used to denote interactions between animals but
not between animals and their inanimate environment (see also Friederichs 1963).
This view thus comes closer to the original meaning of the word “sociology” as
coined in the nineteenth century by Auguste Comte to describe the relations
between human individuals and society. The equivalent to what Braun-Blanquet
described as plant sociology is thus not “animal sociology” but “animal ecology”
in the broadest sense.26
The notion of ecology as a science that studies “wholes” which extend beyond
the biological community – the entire “household of nature” – was developed
mainly by scientists working in aquatic ecology, e.g. August Thienemann and
Richard Woltereck (e.g. Thienemann 1939). The way in which these “wholes” were
conceived, as well as the metaphors used, were highly varied (see Jax 1998; Schwarz
2003 and Chap. 4), even more so after the emergence of systems theories. With the
rise of systems theory in ecology (see Chap. 15) around the middle of the twentieth
century, some authors felt it necessary to modify the traditional – organism-focused
– definitions of ecology and to bring the importance of systems theory and of eco-
systems as the basic objects of ecology to the fore. The web of interrelations and the
emergence of whole systems (including abiotic factors) were given priority here.
23 Braun-Blanquet (1932, p. 1): “Die Pflanzensoziologie, die Lehre von den Pflanzengesellschaften,
auch Vegetationskunde im weitesten Sinn, umfaßt alle das soziale Zusammenleben der Pflanzen
berührenden Erscheinungen”. Braun-Blanquet (1928, p. 1) Instead of providing a translation of
the 1928 text of Braun-Blanquet myself, I am here using the text of the English translation of his
book from 1932.
24 Braun Blanquet (1932, p. 1f): “…das Gesellschaftsgefüge (Organisation oder Struktur)(...), der
Gesellschaftshaushalt (Synökologie)(...), die Gesellschaftsentwicklung (Syngenetik)(...), die
Gesellschaftsverbreitung (Synchorologie) (...) und die Klassifikation oder Systematik der
Pflanzengesellschaften”. Braun-Blanquet (1928, p. 2). This definition of plant sociology was later adopted
by the International Congress of Botany in Paris in 1954 (according to Braun-Blanquet 1964, p. 22).
25 Allaby (1994, p. 302); a similar definition can be found in Calow (1998).
26 In fact the conceptual development of plant and animal ecology, as of aquatic and terrestrial
ecology, has only partially overlapped, but instead, especially in the early years of ecology,
occurred in rather separated circles (see also McIntosh 1985).
167
Thus Eugene Odum remarked: “As you know ecology is often defined as: The
study of interrelationships between organisms and environment. I feel that this
conventional definition is not suitable; it is too vague and too broad. Personally I
prefer to define ecology as: The study of the structure and function of ecosystems.
Or we might say in a less technical way: The study of the structure and function of
nature”. Odum (1962, p. 108). In a similar way, Spanish ecologist Ramon Margalef
defined ecology as the “biology of ecosystems” (Margalef 1968, p. 4).
A more recent view, which attempted to integrate the different aspects described
above, is the definition of ecology given by Gene Likens: “Ecology is the scientific
study of the processes influencing the distribution and abundance of organisms, the
interactions among organisms, and the interaction between organisms and the
transformation and flux of energy and matter” (Likens 1992, p. 8). This definition
represents a continuation of developments in twentieth century ecology: while
focusing on organisms as the core of all ecology (paraphrasing Andrewartha and
Birch 1961; Krebs 198527) it also places emphasis on the “transformation and flux
of energy and matter”, thus alluding to those aspects most frequently seen as
constituting the core of ecosystem ecology.
Another point of controversy regarding the delineation of the ecology concept
should be mentioned again here, having been touched upon only briefly above. This
is the relationship between ecology and biogeography, which was and is perceived in
very different ways and has been discussed extensively time and again (e.g. McMillan
1956; Major 1958; Müller 1980; Browne 2000), not only as a conceptual issue but
also as a matter of marking off (and occupying) professional fields of activity. For
Haeckel both ecology and biogeography (“chorology”) were biological (physiological)
sub-disciplines of equal status (see Chap. 10). As quoted above, Warming (1896, p. 2)
distinguished “ecological plant geography” (i.e. ecology in his sense) from traditional
“floristic (and at the same time historical) plant geography”.28 In doing so, he included
some issues of plant biogeography in the domain of ecology, as did many other
authors (such as H.C. Cowles29). Biogeography is thus frequently understood as a kind
of “border science”, one side belonging to biology, the other to geography (Hesse
1924; Major 1958; Illies 1971; Leser et al. 1993). For some authors, by contrast,
biogeography is an umbrella science, extending far beyond ecology. For these authors,
ecology is a subsidiary branch of biogeography.30 At the same time, biogeography is
12 Stabilizing a Concept
27 Krebs 1985, p. 4: “Ecology is the scientific study of the interactions that determine the distribution
and abundance of organisms”.
28 This approach is similar to that of Krebs (1985, p. 41). The historical dimension of biogeographical
explanations is emphasized by many authors, including Simberloff (1983) and Ricklefs (2004).
29 Cowles (1901, p. 74) made a further distinction between those parts of community-related ecology that
deal with the regional distribution of organisms, which he called “biogeographical ecology”, and those
that deal with local distribution, or local communities, which he called “physiographic ecology”.
30 Dansereau, for example, writes in the preface of his book Biogeography: “The scope of this book
extends across the fields of plant and animal ecology and geography, with many overlaps into
genetics, human geography, anthropology, and the social sciences. All of these together form the
domain of biogeography”. Dansereau (1957, p. v).
168
K. Jax
sometimes perceived as a branch of ecology (this includes, in my view, Walter and
Breckle 1983, even though they do not explicitly use the word biogeography), or
else the two disciplines are considered as being indistinguishable. MacArthur &
Wilson commented in their book Island Biogeography: “Now we both call ourselves
biogeographers and are unable to see any real distinction between biogeography and
ecology”. MacArthur and Wilson (1967, p. v.).
A more recent – albeit not very successful – tendency is the endeavour to create a
terminological distinction between concepts and research approaches of a branch of
“bioecology”31 (as a part of biology) and “geoecology” (as a part of geography), which
are then both seen as constituting the interdisciplinary “umbrella science” of ecology.
This is promoted mainly by Hartmut Leser in Switzerland.32 Whether or not this is a
useful distinction is questionable. Although geographical aspects are important in
ecology, it is always characterized by a concern with living beings and thus is always
an essentially biological discipline. There can be – as Walter and Breckle (1983, p. 1)
remarked – a geography of the moon, but never an ecology of the moon.
More recently, the meaning of “ecology” has been further complicated by its
usage in the environmental movement and within the realm of “political ecology”
(see Chap. 16). Here, ecology has become a buzzword, which is extended to almost
everything (for example, see McIntosh 1985, pp. 6ff). In some quarters it represents
a worldview, while in others it simply stands for “systemic” or for “related to interac-
tions and connections”. However, this is almost exclusively the case when the term
is used outside the narrower scientific community of ecologists.
The meanings of “ecological” and the explicit definitions of “ecology” within
science varied and still vary in the emphasis they give to organism-environment rela-
tions, interactions between organisms, patterns of distribution and abundance, and a
view of the “functioning” of ecological wholes (units) beyond the individual organism.
Struggles over the definition of ecology and its sub-disciplines have thus always been
both about structuring and assessing the complex subject matter of the interdependence
of living and non-living nature as well as about debating the boundaries of institutional
and social groups – in academia and beyond. This process is still ongoing.
31 Walter Taylor used the term “bio-ecology” as far back as 1927. An American textbook from
1939 (Clements and Shelford 1939) also bore this title. In neither case, however, was the term used
as a means of demarcation from geoscience. It was used instead to emphasize the integration of
the two subdisciplines plant ecology and animal ecology, which had commonly been treated in
separate textbooks prior to this, in order to create an ecology of all living beings. The term
“bioecology” has not met with broad acceptance in ecology.
32 Leser (1984, 1991); see also the Diercke Dictionary Ecology and Environment, Leser et al.
(1993); also Rowe and Barnes (1994).
169
References
Allaby M (ed) (1994) The concise Oxford dictionary of ecology. Oxford University Press,
Oxford
Allee WC, Emerson AE, Park O, Park T, Schmidt KP (1949) Principles of animal ecology.
Saunders, Philadelphia
Andrewartha HG, Birch LC (1961) The distribution and abundance of animals. University of
Chicago Press, Chicago
Braun-Blanquet J (1928) Pflanzensoziologie. Springer, Berlin
Braun-Blanquet J (1932) Plant sociology: the study of plant communities. McGraw-Hill, New York
Braun-Blanquet J (1964) Pflanzensoziologie. Grundzüge der Vegetationskunde, 3 edn. Springer,
Berlin, Wien, New York
Browne J (2000). History of biogeography. Encyclopedia of Life Sciences [online]. Wiley &
Blackwell. URL: www.els.net
Calow P (ed) (1998) The encyclopedia of ecology and environmental management. Blackwell,
Oxford
Cittadino E (1980) Ecology and the professionalization of botany in America, 1890-1905. Stud
Hist Biol 4:171–198
Cittadino E (1990) Nature as the laboratory: Darwinian plant ecology in the German Empire,
1880–1900. Cambridge University Press, Cambridge
Clements FE (1905) Research methods in ecology. The University Publishing Company, Lincoln
Clements FE, Shelford VE (1939) Bio-ecology. Wiley, New York
Cowles HC (1901) The physiographic ecology of Chicago and vicinity; a study of the origin,
development, and classification of plant societies. Bot Gaz 31:73–108, 145–182
Cowles HC (1904) The work of the year 1903 in ecology. Science 19:879–885
Dansereau P (1957) Biogeography: an ecological perspective. Ronald Press, New York
Elton C (1927) Animal ecology. Sidgwick & Jackson, London
Flahault C, Schröter C (eds) (1910) Phytogeographische Nomenklatur. III. Internationaler
Botanischer Kongress, Brüssel 1910. Zürcher & Furrer, Zürich
Friederichs K (1963) Über den Gebrauch der Worte und Begriffe “Gesellschaft” und “Soziologie”
in verschiedenen Sparten der Wissenschaft. KZfSS 15:449–461
Gleason HA (1917) The structure and development of the plant association. Bull Torrey Bot Club
44:463–481
Gleason HA (1926) The individualistic concept of the plant association. Bull Torrey Bot Club
53:7–26
Grisebach (1838) Über den Einfluß des Klimas auf die Begrenzung der natürlichen Floren. In:
Grisebach A (ed) Gesammelte Abhandlungen und kleinere Schriften zur Pflanzengeographie.
Verlag von Wilhelm Engelmann, Leipzig, pp 1–29
Haberlandt G (1884) Physiologische Pflanzenanatomie. Engelmann, Leipzig
Hagen JB (1988) Organism and environment: Frederic Clements´s vision of a unified physiologi-
cal ecology. In: Rainger, Ronald, Benson KR, Jane Maienschein (eds) The American develop-
ment of biology. University of Pennsylvania Press, Philadelphia, pp 257–280
Hagen JB, (1986) Ecologists and taxonomists: divergent traditions in twentieth-century plant
geography. J Hist Biol 19:197–214
Hesse R (1924) Tiergeographie auf ökologischer Grundlage. Gustav Fischer, Jena
Hesse R (1927) Die Ökologie der Tiere, ihre Wege und Ziele. Naturwissenschaften 15:942–946
Alexander von Humboldt (1807) Ideen zu einer Geographie der Pflanzen. In: Dittrich M (1957)
(ed) Ideen zu einer Geographie der Pflanzen, Akademische Verlagsgesellschaft Geest &
Portig, Leipzig, pp 29–50
Illies J (1971) Einführung in die Tiergeographie. Gustav Fischer, Stuttgart
Jax K (1998) Holocoen and ecosystem: on the origin and historical consequences of two concepts.
J Hist Biol 31:113–142
Jax K (2006) The units of ecology: definitions and application. Q Rev Biol 81:237–258
12 Stabilizing a Concept
170 K. Jax
Kohler RE (2002) Landscapes and labscapes: exploring the lab-field border in biology. University
of Chicago Press, Chicago/London
Krebs CJ (1985) Ecology: the experimental analysis of distribution and abundance. Harper &
Row, New York
Leser H (1984) Zum Ökologie-, Ökosystem- und Ökotopbegriff. Nat Land 59:351–357
Leser H (1991) Ökologie wozu? Der graue Regenbogen oder Ökologie ohne Natur. Springer,
Berlin
Leser H, Streit B, Haas H-D, Huber-Fröhli J, Mosimann T, Paesler R (1993) Diercke Wörterbuch
Ökologie und Umwelt. Band 2 N-Z. dtv/Westermann, München, Braunschweig
Likens GE (1992) The ecosystem approach: its use and abuse. Ecology Institute, Oldendorf/Luhe
MacArthur RH, Wilson EO (1967) The theory of island biogeography. Princeton University Press,
Princeton
Madison Botanical Congress (1894) Proceedings of the Madison Botanical Congress, Madison,
23–24 Aug 1893
Major J (1958) Plant ecology as a branch of botany. Ecology 38:352–363
Margalef R (1968) Perspectives in ecological theory. The University of Chicago Press, Chicago/
London
McIntosh RP (1985) The background of ecology: concept and theory. Cambridge University
Press, Cambridge
McMillan C (1956) The status of plant ecology and plant geography. Ecology 37:600–602
Müller P (1980) Biogeographie. Eugen Ulmer, Stuttgart
Nichols GE (1923) A working basis for the ecological classification of plant communities.
Ecology 4:11–23, 154–179
Odum EP (1962) Relationship between structure and function in the ecosystem. Jpn J Ecol
12:108–118
Ricklefs RE (2004) A comprehensive framework for global patterns in biodiversity. Ecol Lett
7:1–15
Rowe JS, Barnes BV (1994) Geo-ecosystems and bio-ecosystems. Bull Ecol Soc Am 75:40–41
Schimper AFW (1898) Pflanzengeographie auf physiologischer Grundlage. Gustav Fischer, Jena
Schwarz AE (2003) Wasserwüste - Mikrokosmos - Ökosystem. Eine Geschichte der “Eroberung”
des Wasserraums. Rombach-Verlag, Freiburg
Semper K (1868) Reisen im Archipel der Phillipinen. Zweiter Teil: Wissenschaftliche Resultate.
Erster Band: Holothurien. Verlag von Wilhelm Engelmann, Leipzig
Semper K (1880) Die natürlichen Existenzbedingungen der Thiere. Brockhaus, Leipzig
Semper K (1881) Animal life as affected by the natural conditions of existence. D. Appleton, New York
Shelford VE (1913) Animal communities in temperate America. University of Chicago Press,
Chicago
Simberloff DS (1980) A succession of paradigms in ecology: essentialism to materialism and
probabilism. Synthese 43:3–39
Simberloff DS (1983) Biogeography: the unification and maturation of a science. In: Brush AH, Jr
Clark GA (eds) Perspectives in ornithology. Cambridge University Press, Cambridge, pp 411–455
Tansley AG (1914) Presidential address to the first annual general meeting of the British ecologi-
cal society. J Ecol 2:194–202
Tansley AG (1920) The classification of vegetation and the concept of development. J Ecol
8:118–149
Taylor WP (1927) Ecology or bio-ecology. Ecology 8:280–281
Thienemann (1939) Grundzüge einer allgemeinen Ökologie. Arch Hydrobiol 35:267–285
Trepl L (1987) Geschichte der Ökologie. Vom 17. Jahrhundert bis zur Gegenwart. Athenäum,
Frankfurt/M
Walter H, Breckle SW (1983) Ökologie der Erde. Ökologische Grundlagen in globaler Sicht, vol 1.
Gustav Fischer, Stuttgart
Warming E (1895) Plantesamfund. Grundtræk af den økologiske plantegeografi. Philipsen,
Kjobenhavn
Warming E (1896) Lehrbuch der ökologischen Pflanzengeographie: Eine Einführung in die
Kenntnis der Pflanzenvereine. Gebrüder Bornträger, Berlin
171
An important step in consolidating ecology as a “self conscious” science was
the formation of scientific societies explicitly devoted to ecology or particular parts
of it.
The first ecological societies were formed in Great Britain (1913) and the USA
(1915). From the outset, these societies intended to accommodate both botanists and
zoologists, although this did not mean that close co-operation between the two fields
became common in practice. The forerunner of the British Ecological Society (BES)
was the British Vegetation Committee, initiated by Arthur Tansley in 1904, a body
established with the primary purpose of surveying and mapping Britain’s vegetation.1
The BES also published the Journal of Ecology which, like the society as a whole,
was initially dominated by plant ecologists. Two decades later (in 1932) the society
established a second journal on the initiative of Charles Elton (Journal of Animal
Ecology), devoted to zoological and ecological studies. Two years after the founding
of the BES, the Ecological Society of America (ESA) came into being along with
its new journal “Ecology”.2 (see Part VI, this volume for further details of local
traditions).3
In contrast to specialised branches of ecology and related research fields, inter-
national societies covering ecology as a whole were much less successful and are a
rather late development. The International Association for Ecology (INTECOL)
was founded in 1967 as part of the Section of the Environment of the International
Union of Biological Sciences (IUBS), and has held several major conferences (one
every 3 years since 1975). However, it has never achieved the renown of some of
1 For the special role of this committee, see Fischedick (2000).
2 The journal itself started in 1920, i.e. five years after the founding of the society.
3 For a history of the British Ecological Society, see Salisbury (1964) and Sheail (1987), for that
of the Ecological Society of America, Burgess (1977).
K. Jax ()
Department of Conservation Biology, Helmholtz Centre for Environmental Research (UFZ),
Permoserstr. 15, 04318 Leipzig, Germany
e-mail: kurt.jax@ufz.de
Chapter 13
Formation of Scientific Societies
Kurt Jax
A. Schwarz and K. Jax (eds.), Ecology Revisited: Reflecting on Concepts, Advancing Science,
DOI 10.1007/978-90-481-9744-6_13, © Springer Science+Business Media B.V. 2011
172
the national societies or more specialised global organisations. At the European
level, there is a Federation of Ecological Societies (European Ecological Federation,
EFF), but no distinct Society in its own right.
Within the more specialised branches of ecology, the most successful interna-
tional societies have been formed within aquatic ecology and vegetation science.
The special role played within ecology by aquatic ecology is demonstrated by the
fact that separate societies were formed for these disciplines. At the international
level, the International Limnological Society (SIL) was founded by August
Thienemann and Einar Naumann in 1922 (see Elster 1974; Rodhe 1974; Steleanu
1989; Schwarz 2003) and became a major focal point for research on freshwater ecology.
In some countries, such as the USA,4 national societies for limnology and/or marine
ecology (and, even more broadly, oceanography) also developed, though mostly not
until the second half of the twentieth century. Limnology and marine ecology were
among those parts of ecology that were able to establish themselves rather early on
in university departments and other institutions, not least because these fields were
often closely related to fisheries and water treatment.5 Aquatic research, especially in
marine research societies and institutions, was and is generally much broader in scope
than “mere” marine ecology or freshwater ecology, i.e. it also involves much hydro-
graphic research.6 A combined emphasis on or even the study of both fields (marine
and freshwater ecology) in scientific societies, as in the American Society of
Limnology and Oceanography, remains a very rare exception, however.
Focusing on theoretical and practical studies of vegetation, the International
Association for Vegetation Science (IAVS) was founded in 1947 in Hilversum/
Netherlands. Its precursor was the (very short-lived, on account of World War II)
International Phytosociological Society (IPS) created in 1939 with its headquarters
in Montpellier/France. In fact, the IPS did not consider itself as being “only”
devoted to ecological research. In the founding statement of the society, as
published in the journal Ecology (Anonymous 1939, p. 110) the first aim of the
society is stated as: “The development of phytosociology (and geobotany) by a
closer collaboration between phytosociologists and ecologists”, thus identifying
phytosociology and (plant) ecology as overlapping but not identical fields.
No similar organisation exists in relation to animal ecology. In contrast to
aquatic ecology, the establishment of academic institutions was much more difficult
K. Jax
4 The Limnological Society of America was formed in 1936 and in 1948 merged with the
Oceanographic Society of the Pacific to form the American Society of Limnology and
Oceanography (ASLO).
5 It is for this reason that marine biological stations and several national marine research commis-
sions and associations, such as the Marine Biological Association of the United Kingdom, devel-
oped as early as the late nineteenth century (Hedgpeth 1957).
6 The founders of the SIL emphasized that limnology is “the science of freshwater as a whole and
includes everything that concerns freshwater (…), freshwater hydrography and freshwater biology”.
(“die Wissenschaft vom Süßwasser im ganzen umfaßt alles, was das Süßwasser betrifft (…), limnische
Hydrographie und limnische Biologie”) (Naumann and Thienemann 1922, p. 585; emphasis in
original).
173
for terrestrial ecology.7 This changed dramatically along with the increasing awareness
of environmental problems from the 1960s onwards, with the creation of many new
chairs in ecology and ecology departments.
References
Anonymous (1939) The international society of phytosociology. Ecology 20:110
Burgess RL (1977) The ecological society of America. Historical data and some preliminary
analyses. In: Egerton FN (ed) History of American ecology. Arno Press, New York, pp 1–24
Crowcroft P (1991) Elton’s ecologists. A history of the bureau of animal population. Chicago
University Press, Chicago
Elster H-J (1974) History of limnology. Mitteilungen. Internationale Vereinigung für Limnologie
20:7–30
Fischedick KS (2000) From survey to ecology: the role of the British Vegetation Committee,
1904–1913. J Hist Biol 33:291–314
Godwin H (1977) Sir Arthur Tansley: the man and the Subject. J Ecol 65:1–26
Hedgpeth JW (1957) Introduction. Treatise on marine ecology and paleoecology. Geol Soc Am
Mem 67:1–16
Naumann E, Thienemann A (1922) Vorschlag zur Gründung einer internationalen Vereinigung für
theoretische und angewandte Limnologie. Arch Hydrobiol 13:585–605
Rodhe W (1974) The International Association of Limnology: creation and functions. Mitteilungen.
Internationale Vereinigung für Limnologie 20:44–70
Salisbury E (1964) The origin and early years of the British Ecological Society. J Ecol
52(Suppl):13–18, Appendix p. 244
Schwarz AE (2003) Wasserwüste – Mikrokosmos - Ökosystem. Eine Geschichte der “Eroberung”
des Wasserraums. Rombach-Verlag, Freiburg
Sheail J (1987) Seventy-five years in ecology. The British Ecological Society. Blackwell, Oxford
Steleanu A (1989) Geschichte der Limnologie und ihrer Grundlagen. Haag & Herchen, Frankfurt/
Main
13 Formation of Scientific Societies
7 The fate of the two most eminent British ecologists of the first half of the twentieth century,
Charles Elton and Arthur Tansley, is a good example for these problems: Elton struggled –
successfully – to establish a kind of semi-private institution, the Bureau of Animal Population at
Oxford University, which, however, did not survive after his retirement (Crowcroft 1991). Tansley,
one of the founders of the BES and an internationally renowned plant ecologist and Fellow of the
Royal Society, only received a chair at Oxford at the age of 56 (in 1927; see Godwin 1977).
175
The conceptual foundations of ecology were developed rather independently in
different biological fields (McIntosh 1985; Jax 2000; Schwarz 2003), leading early
on to an array of subdivisions within the emerging discipline. These subdivisions
result in part from research traditions that go back beyond the formation of ecology
as a science and in part from new distinctions arising out of specialisations and new
emerging topics within ecology. One distinction that was especially important in
shaping the character of ecology as a concept was that created between those
scientific fields within ecology that dealt with individual organisms and those that
dealt with groups of organisms, in particular the distinction between autecology
and synecology and, later on, population ecology.
The first kind of subdivisions mentioned above refer to the division into fields
dealing with particular taxonomic groups, namely animal and plant ecology (more
recently also microbial ecology), and those focusing on the differences between
different types of environments (terrestrial, freshwater, marine).
Plant and animal ecology as well as aquatic and terrestrial ecology at first developed
quite independently from each other. Even more distinct fields were paleoecology
and parasitology.1 The existence of these separate branches of ecology was in part
brought about by the traditional academic divisions of biology into, for example,
Chapter 14
The Fundamental Subdivisions of Ecology
Kurt Jax and Astrid Schwarz
1 Parasitology is generally addressed in the context of disease research in human and veterinary
medicine or plant pathology. The term “paleoecology” (or “paleo-ecology”) was coined by
Frederic Clements (1916, p. 279), while studies on the relationship between fossil organisms and
their environment had already begun to be conducted by Edward Forbes and others by at least
1840 (see Cloud 1959, p. 927f; Hecker 1965, pp. 1ff; McIntosh 1985, pp. 98ff). Paleoecology
often remained equally or even more closely related to geology than to biology and ecology.
K. Jax (*)
Department of Conservation Biology, Helmholtz Centre for Environmental Research (UFZ),
Permoserstr. 15, 04318 Leipzig, Germany
e-mail: kurt.jax@ufz.de
A. Schwarz
Institute of Philosophy, Technische Universität Darmstadt, Schloss, 64283 Darmstadt, Germany
e-mail: schwarz@phil.tu-darmstadt.de
A. Schwarz and K. Jax (eds.), Ecology Revisited: Reflecting on Concepts, Advancing Science,
DOI 10.1007/978-90-481-9744-6_14, © Springer Science+Business Media B.V. 2011
176
K. Jax and A. Schwarz
zoology and botany, but also by the specific characteristics of their objects and the
different methods required for their investigation.
Botany (and plant geography), zoology, and the aquatic sciences (marine as well
as freshwater) were the traditional fields in which ecology first developed. Plant
ecology was the forerunner in shaping ecology and contributed crucially to its forma-
tion as a self-conscious science – an issue that was also acknowledged by zoologists
(e.g. Shelford 1915, p. 1; Hesse 1927). A major step in the stabilisation of ecology
as a distinct research field with its own concepts was the merging of the biogeographic
and the physiological perspective on plants and, especially, plant communities, which
occurred during the last decade of the nineteenth century (see Chap. 12). Animal
ecology, by contrast, remained a science without a strong common conceptual core
or a common sense of constituting a scientific community in its own right for at
least another 2 decades (Trepl 1987, p. 160f).
In terms of their training and the objects they studied, aquatic ecologists were
mainly zoologists. As with plant ecology, limnology and oceanography had their
roots in a longer tradition of geographical research on seas and freshwater bodies
(in particular lakes), which were systematically related to biological studies only
towards the end of the nineteenth century. Aquatic ecologists were much more
inclined than terrestrial ecologists to emphasize and investigate the “whole” web of
interrelations in a particular place (e.g. a lake) and thus to consider larger wholes,
which included the physical and chemical properties of the habitat they studied, as
the basic objects of ecology.2
Beyond the question of which kinds of organisms constituted the object of study,
another important subdivision was created by the delimitation of those scientific
branches within biology (and thus also within ecology) that dealt with individual
organisms from those that dealt with groups of organisms. Carl Schröter and Oskar
Kirchner in 1902 distinguished between the ecology of individual organisms, or
autecology (German: Autökologie), and that of species assemblages, or synecology
(German: Synökologie). The term Synökologie was coined thus: “I propose to intro-
duce for this important discpline [the science of plant communities] the name ‘forma-
tion science’ or ‘synecology’, from = with, and oikos = house, i.e., the science of the
plants that live together, and at the same time the science of the plants that seek out
analogous ecological conditions”.3 A similar division of ecology into two “phases”
– as he called it – was also undertaken by Henry Chandler Cowles, although he did
not provide specific names for these subdisciplines: “Whatever its limits may be,
ecology is essentially a study of origins and life histories, having two well-marked
phases; one phase is concerned with the origin and development of plant structures,
2 See Schwarz (2003) for a history of the development of aquatic ecology.
3 “Ich schlage vor, für diese wichtige Disziplin [die Lehre von den Pflanzengesellschaften] den
Namen ‚Formationslehre‘ oder ‚Synökologie‘ einzuführen, von συν=mit, zusammen wohnen und
oikos = Haus, also die Lehre von den Pflanzen, welche zusammen wohnen, und zugleich die
Lehre von den Pflanzen, welche analoge ökologische Bedingungen aufsuchen” (Schröter and
Kirchner 1902, p. 63).
177
14 The Fundamental Subdivisions of Ecology
the other with the origin and development of plant societies or formations” (Cowles
1901, p. 73). Other authors substituted the words Biocoenotik (biocoenology) (Gams
1918) or Biosoziologie (biosociology) (Du Rietz 1921) for “synecology”. This
distinction, between the biology and ecology of individual organisms and that of
groups of organisms, was emphasized in particular by Gams (1918)4 and Schwenke
(1953) and considered by them as being of greater significance than the distinctions,
for example, between ecology and chorology, or physiology and morphology.5
The division of ecology into autecology, population ecology (also: demecology,
in German: Demökologie), and synecology, which is currently common, is of more
recent origin.6 Population ecology is sometimes subsumed under synecology and
sometimes considered a (sub)discipline in its own right.
Population ecology – despite also having far-reaching roots, especially in human
demography (see e.g. Hutchinson 1978, pp. 5ff) – made a rather late appearance as a
subdiscipline of ecology. Its growth, though rapid, only began to occur during the
1920s and 1930s. Despite its late appearance on the scene, population ecology was
the first subdiscipline of ecology to enter the stage of mathematical theory building
(Kingsland 1985/1995). Alongside autecology, population ecology was that part of
ecology which pursued most stringently the “hard science” approach. Although it was
claimed (by Thomas Park (1946), for example) that population ecology paid no heed
to boundaries of habitat and taxonomy in the scope of its studies, it was in fact almost
completely a zoological enterprise. Even though Clements and Tansley also devoted
time to working experimentally on plant populations, a population ecology of plants
was practically non-existent in the first half of the twentieth century and was system-
atically developed only during the 1960s and 1970s in Britain by John Harper.7
The investigation of “ecosystems” is most commonly subsumed under synecology.
While it is common in English speaking countries to distinguish between population
ecology, community ecology and ecosystem ecology – especially after Eugene Odum
and his highly influential textbook8 – the subdivision of ecology based on “individuals,
populations and communities”9 found in the most popular contemporary English
textbook of ecology (Begon et al. 1990, 1996, 2005) – is again close to that used for
4 For Gams (1918, p. 297), this distinction was a fundamental subdivision of the whole field of
biology, which he divided into the two major branches of Idiobiologie (idiobiology: the science of
individual organisms) and Synbiologie (synbiology: the science of biological communities).
5 See especially van der Klaauw (1936) for a detailed discussion of this subdivision.
6 We managed to find this systematic division in the German-language literature first in
Schwerdtfeger 1968, who apparently coined the term Demökologie as an alternative word for “popu-
lation ecology”. Schwerdtfeger himself points out that population ecology had already been estab-
lished previously as a separate branch between autecology and synecology in the English-language
literature; see, e.g. Park (1946).
7 The seminal textbook of plant population biology was Harper (1977).
8 First edition 1953; revised and extended editions followed in 1959 and 1971.
9 In the 4th edition of the book (2005) the part formerly headed “Communities” is now entitled
“Communities and Ecosystems”.
178
K. Jax and A. Schwarz
a long time in German speaking ecological circles. Depending on author and view-
point one the “levels” is sometimes omitted in one or the other textbook. A uniform
scheme does not exist here. More recently, other authors (e.g. Jones and Lawton
1995) emphasize the distinction between an ecology of populations and communi-
ties on the one hand (focusing on specific species’ interactions) and ecosystem ecol-
ogy on the other, the latter seen as dealing mainly with flows of energy and matter.
During the 1980s, landscape ecology became yet another subdiscipline of ecology, for
the most part emphasizing large-scale topological aspects of ecological phenomena.10
Distinctions vis-à-vis the other subdisciplines mentioned above, especially ecosystem
ecology, are not always clear-cut.
Since the 1970s and 1980s, when ecology became a more established and popular
science, numerous additional branches of ecology have been constituted, each one
focusing on specific methods, perspectives, and objects. They include, for example,
behavioral ecology, molecular ecology, functional ecology, microbial ecology, and
evolutionary ecology. Some of these new ecological specialisms and their scientific
communities overlap with the broader, more traditional branches, while others
(such as microbial ecology) are addressed by rather distinct and separate research
communities.
References
Begon M, Harper JL, Townsend CR (1990) Ecology: individuals, populations and communities,
2nd edn. Blackwell, Oxford
Begon M, Harper JL, Townsend CR (1996) Ecology. Individuals, populations and communities,
3 edn. Blackwell, Oxford
Begon M, Townsend CR, Harper JL (2005) Ecology: from individuals to ecosystems, 4th edn.
Blackwell, Oxford
Clements FE (1916) Plant succession: an analysis of the development of vegetation. Carnegie
Institution of Washington, Washington, DC, Publication No. 242
Cloud PE Jr (1959) Paleoecology: retrospect and prospect. J Paleontol 33:926–962
Cowles HC (1901) The physiographic ecology of Chicago and vicinity; a study of the origin,
development, and classification of plant societies. Bot Gaz 31:73–108, 145–182
Du Rietz E G (1921) Zur methodologischen Grundlage der modernen Pflanzensoziologie. Adolf
Holzhausen, Wien
Gams H (1918) Prinzipienfragen der Vegetationsforschung. Ein Beitrag zur Begriffsklärung und
Methodik der Biocoenologie. Vierteljahresschrift der Naturforschenden Gesellschaft in Zürich
63:293–493
10 In contrast to Anglo-Saxon tradition, landscape has been an object of German ecology already
since the first half of the twentieth century. However, these traditions derived from a different
context, connecting cultural and natural aspects of human living spaces, and not from a purely
scientific approach (see Trepl 1995; Klink et al. 2002, and Chap. 25).
179
14 The Fundamental Subdivisions of Ecology
Harper JL (1977) Population biology of plants. Academic Press, London
Hesse R (1927) Die Ökologie der Tiere, ihre Wege und Ziele. Naturwissenschaften 15:942–946
Hecker [Gekker] RF (1965) Introduction to paleoecology, American Elsevier, New York
Hutchinson GE (1978) An introduction to population ecology. Yale University Press, New Haven/
London
Jax K (2000) History of ecology. In: Encyclopedia of Life Sciences. [online]. Wiley. URL: www.
els.net
Jones CG, Lawton JH (eds) (1995) Linking species and ecosystems. Chapman & Hall, New York
Kingsland SE (1985/1995) Modeling nature: episodes in the history of population ecology.
University of Chicago Press, Chicago
van der Klaauw CJ (1936) Zur Aufteilung der Ökologie in Autökologie und Synökologie, im Lichte
der Ideen als Grundlage der Systematik der zoologischen Disziplinen. Acta Biotheor 2:195–241
Klink H-J, Marion P, Bärbel T, Gunther T, Martin V, Uta S (2002) Landscape and landscape ecol-
ogy. In: Bastian O, Uta S (eds) Developments and perspectives of landscape ecology. Kluwer,
Dordrecht, pp 1–47
McIntosh RP (1985) The background of ecology: concept and theory. Cambridge University
Press, Cambridge
Odum EP (1953) Fundamentals of ecology, 1st edn. W.B. Saunders, Philadelphia
Park T (1946) Some observations on the history and scope of population ecology. Ecol Monogr
16:313–320
Schröter Carl, Oskar Kirchner (1902) Die Vegetation des Bodensees, 2. Teil. Lindau i. B.:
Kommissionsverlag der Schriften des Vereins der Geschichte des Bodensees und seiner
Umgebung von Joh. Thom. Stettner
Schwarz AE (2003) Wasserwüste – Mikrokosmos – Ökosystem. Eine Geschichte der “Eroberung”
des Wasserraums. Rombach-Verlag, Freiburg
Schwenke W 1953. Biozönotik und angewandte Entomologie. Beiträge zur Entomologie 3,
Beiheft: 86–162
Schwerdtfeger F (1968) Ökologie der Tiere. Band II: Demökologie: Struktur und Dynamik
tierischer Populationen. Paul Parey, Hamburg/Berlin
Shelford VE (1915) Principles and problems of ecology as illustrated by animals. J Ecol 3:1–23
Trepl L (1987) Geschichte der Ökologie. Vom 17. Jahrhundert bis zur Gegenwart. Athenäum,
Frankfurt/Main
Trepl L (1995) Die Landschaft und die Wissenschaft. In: Erdmann K-H, Kastenholz HG (eds)
Umwelt- und Naturschutz am Ende des 20, Jahrhunderts. Probleme, Aufgaben und Lösungen.
Springer, Berlin/Heidelberg/New York, pp 11–26
Part V
“Ecology”, Society and the Systems View
in the Twentieth and Twenty-first Century
183
Chapter 15
The Rise of Systems Theory in Ecology
Annette Voigt
The emergence of systems theory in ecology, particularly during the 1950s and
1960s, was accompanied by the hope that ecology might turn into an exact science
with prognostic potential and a set of uniform theoretical foundations. The impact
of systems theory on ecology was manifested mainly in the formulation and devel-
opment of ecosystem theory. The widely-held view is that ecosystem theory is
concerned primarily with units comprising communities of organisms of various
species and the abiotic environment of these communities. The components of
systems are seen to interact with one another.
The main elements in any historical reconstruction of the emergence of ecosystem
theory include the establishment of general systems theory and its associated theories
(including cybernetics, information theory etc.), the introduction of the term
“ecosystem” and early ecosystem theories.
General Systems Movement
During the 1940s a variety of approaches were developed in different parts of
Europe, the USA and the USSR, which were later to become united under the
rubric of “systems theory”. Most of these theories and practices came about as a
result of encounters between scientists from different disciplines. Perhaps the one
motivation they can all be said to have shared was an interest in the scientific
description of “gestalt”, a term found throughout the scientific literature of the first
few decades of the twentieth century, in a wide range of disciplines.1 Otherwise,
1 The most prominent is probably the gestalt concept in psychology (Köhler 1920), but see also
the field concept(s) in physics, and the “gestalt laws” (Bertalanffy 1926, 1929). The role of the
latter with respect to ecology is discussed in Schwarz (1996).
A. Voigt (*)
Urban and Landscape Ecology Group, University of Salzburg, Hellbrunnerstraße 34,
5020 Salzburg, Austria
e-mail: annette.voigt@sbg.ac.at
A. Schwarz and K. Jax (eds.), Ecology Revisited: Reflecting on Concepts, Advancing Science,
DOI 10.1007/978-90-481-9744-6_15, © Springer Science+Business Media B.V. 2011
184
A. Voigt
these approaches all differ in terms of their aims, their problem focus and their
institutional background. Approaches with more of an engineering, mathematical
or physical background include cybernetics (Wiener 1948) and information theory
(Shannon and Weaver 1949); other theories, such as game theory (Neumann and
Morgenstern 1944), operations research (Churchman et al. 1957), action theory
(Parsons 1937) and general systems theory (GST), emphasize more psychological,
physiological and philosophical aspects.
The major figures regarded as the founders of GST are economist Kenneth
E. Boulding, neurophysiologist Ralph W. Gerard, biomathematician Anatol
Rapoport and, in particular, biologist Ludwig von Bertalanffy.2 Bertalanffy started
out from the problem of how to provide a scientific explanation of “life”. His theory
of the organism as a hierarchically organised, open system was intended in part as
a means of overcoming the dispute in biology between mechanism and vitalism
(Bertalanffy 1932, 1949). Thus, organisms were to be described in non-reductionist
terms, as living wholes, yet still within a scientific framework. Taking this organismic
systems theory as a starting point, and having observed certain parallels between
the concepts and models found in different scientific disciplines, Bertalanffy, from
the 1950s onwards, began to define a generalised systems concept, related to all
“sets of elements standing in interrelation”3: technical systems, organisms, social
systems, and so on.4 According to Bertalanffy GST is a new scientific logico-
mathematical discipline: ‘Its subject matter is formulation of principles that are
valid for ‘systems’ in general, whatever the nature of their component elements and
the relation or ‘forces’ between them’ (Bertalanffy 1968, p. 36).
The new thing about GST that set it apart from prevailing understandings of
science up until then was that its aim was to reach a “new level” of theory making:
“Developing unifying principles running ‘vertically’ through the universe of the
individual sciences, this theory brings us nearer to the goal of the unity of science”
(Bertalanffy 1968, p. 37). As such, it ignored the division between the human and
the natural sciences, which had become institutionalised in the nineteenth century,
and applied mathematical procedures to areas where, up until then, hermeneutical
methods had seemed appropriate (Lilienfeld 1978; Müller 1996). The aim, how-
ever, was not to seek explanations that reduced phenomena to their elemental
“fundamental units” (molecules and so on), but rather to provide a mathematical
description of the “system as a whole”, one in which the system’s elements were to
be regarded primarily from the point of view of their function in relation to other
elements or to “the whole”. Approaches adopted within information theory and
cybernetics in the 1950s extended systems theory by adding an explicitly social
scientific dimension, since it is possible to apply a mathematical concept of
2 Boulding 1941, 1953, 1956; Bertalanffy 1950, 1951, 1955; Gerard 1940, 1953; Rapoport 1947,
1950. Also cf. Davidson 1983; Müller 1996; Hammond 2003.
3 “A system can be defined as a complex of interacting elements p1, p2 … pn. Interaction means
that the elements stand in a certain relation, R, so that their behaviour in R is different from their
behaviour in another relation, R¢.” (Bertalanffy 1950, p. 143).
4 Bertalanffy 1950, 1955, 1968; also cf. Müller 1996; Schwarz 1996; Voigt 2001.
185
15 The Rise of Systems Theory in Ecology
information to the sphere of the social without opening oneself up to the accusation
of naturalisation. In addition, cybernetic concepts enable a connection to be made
to mathematical constructions and technological issues.
“Systems theory” thus encompasses various approaches that have proven their
worth in describing, monitoring and constructing complex systems.5 Systems are
classified according to their characteristics, while the specific focus may be on their
relationships to their environment, their complexity, their mode of self-organisation
or the capacity of a system for feedback. Systems theoretical approaches exist
today in all those sciences that see their objects of interest as having, at least in part,
a systemic character; these include – to name just a few – sociology, psychology,
geography, physics, cognitive and neurosciences and ecology.
Historical Overview of Early Ecosystem Theory6
Ecology developed its own systemic theories early on, later applying systems theo-
retical approaches of different kinds.7
In 1935 vegetation ecologist A.G. Tansley introduced the term “ecosystem” as a
“fundamental concept in ecology”. Tansley’s new concept represented a response
to the debate about the structure and organisational form of units of vegetation in
ecology.8 On one side of this debate was the holistic-organicist approach (e.g.
Clements 1916, 1936; Friederichs 1927, 1934, 1937; Phillips 1934, 1935; Clements
and Shelford 1939), according to which a biotic community (“Lebensgemeinschaft”)
is essentially determined by internal, functional relationships of dependency
between individual organisms. As a whole, it has the character of a superorganism
and is usually conceived of as a real unit. At the opposite end of the debate was the
reductionist-individualist approach (e.g. Gleason 1917, 1926; Gams 1918;
Ramensky 1926), which held that the term “association” relates to temporary com-
binations of species determined both by needs that either coincide or are comple-
mentary and by the random character of immigration. It is accepted that this
“association” is only of heuristic use; no real, natural units exist above the level of
the organism.9 Tansley’s ecosystem is neither a superorganism nor a chance
5 More modern systems theoretical variants include non-equilibrium thermodynamics (Prigogine
1955) and theories about adaptive, self-organised and self-referential systems, e.g. autopoiesis
(Maturana and Varela 1987).
6 Hagen (1992) and Golley (1993) provide a detailed account of the history of ecosystem theory.
Also cf. McIntosh 1985.
7 On the theory of the transfer of ideas from ecology to systems theory, Chap. 27.
8 On this debate, see e.g. Tobey 1981, p. 76–109; Worster 1985, p. 205–220; McIntosh 1995,
p. 76–85, 1995; Trepl 1987, p. 139–158; Hagen 1992, p. 15–32; Golley 1993, p. 8–34; Botkin
1990; Jax 2002; Chaps. 19 and 20.
9 The radical individualist position (Peus 1954) rejects the notion of associations as an object of
science because it sees them as “fictions”.
186
A. Voigt
combination: it contains not only organisms, but also their environment,10 and these
components and the interactions that exist between them are viewed in physical
terms. Tansley’s formulation, “The whole method of science […] is to isolate systems
mentally for the purposes of study […]” (1935, p. 299f.) suggests the view that the
system being studied by the scientist is not a real object but rather an idealisation.
No claim is made to be studying all the variables of a phenomenon; instead, abstrac-
tions are made only in relation to those issues that are of interest to the scientist.
Even though the distinction between living and non-living parts is secondary in
Tansley’s system concept, he does refer explicitly to ecological objects and does not
pursue the high level of abstraction found in GST. Tansley distanced himself from
Bertalanffy’s systems theory and, while not doing ecosystem research in today’s
sense (Golley 1993, p. 34), his concept of ecosystem nonetheless contributed
greatly to a physically oriented ecosystem theory.
The period after 1935 saw, on the one hand, the rise of theories oriented towards
physics, which described the transfer of matter, energy and information within the
ecosystem and, on the other, more biologically-oriented positions, which saw the
key to understanding ecosystems in the characteristics of the populations and indi-
viduals that constitute them (e.g. Lamotte and Bourliere 1978).11
The ecosystem concept was first applied in 1942 by the limnologist Raymond
L. Lindeman. He described a lake as an energetically open ecosystem consisting of
biotic and abiotic components. Organisms are significant only insofar as they fulfil
specific functions within the system. They are arranged in trophic levels (producers,
primary and secondary consumers, decomposers). “The basic process in trophic
dynamics is the transfer of energy from one part of the ecosystem to another”.
(Lindeman 1942, p. 400). Energy from the sun is accumulated in the producers by
means of photosynthesis, and only a portion of this energy is transferred via con-
sumption to the next level – everything else is lost (by respiration and decomposi-
tion). It is possible to quantify both the productivity of each level and the degree of
efficiency of the energy transfer, as well as that of both changes in the succession.
This approach enables ecosystems to be described in thermodynamic terms.
One key figure in the history of systems theory in ecology is Lindeman’s teacher
George Evelyn Hutchinson. Hutchinson, an ecologist, was part of the core group
that organised the ten “Macy Conferences” (1946–1953) to explore the possibility
of using scientific ideas that had emerged in the war years as a basis for both inter-
disciplinary research alliances and solving peacefully the complex problems facing
10 Previous approaches to conceptualising communities along with their environment include
Thienemann’s concept of the biosystem (Thienemann and Kieffer 1916) and Friederichs’ concept
of the holocoen (1927). However, these differ from the concept of ecosystem on account of their
holistic-morphological and/or holistic-organicist orientation (Chap. 4).
11 In addition, the concept of ecosystem research is related to the fact that everything is considered
that is relevant ecologically in a specific site. That is, not only all the organisms are considered but
all edaphic and climatic factors as well.
187
15 The Rise of Systems Theory in Ecology
the postwar world.12 The first meeting of this “Cybernetics Group” was called
“Feedback Mechanisms and Circular Causal Systems in Biological and Social
Systems”. It was here, where the different kinds of systems theories discussed
above finally found their way into ecology. With his paper on “Circular Causal
Systems in Ecology” Hutchinson presented a theory for describing a community
using the cybernetic terms feedback and circular causality (Hutchinson 1948).
Within certain boundaries, ecosystems are “self-correcting” by means of “circular
causal paths”, so that conditions of equilibrium prevail. The assumption of regulating
feedback systems forms the basis of both his biogeochemical approach (following
V. Vernadsky) – in which the transfer of substances through the systems is described
quantitatively and without any specifically biological terms – and of his biodemo-
graphic approach, which describes population developments with reference to
quantitative theories of ecology (e.g. Lotka 1925; Volterra 1926). Abiotic and biotic
factors alike are looked at from the point of view of the extent to which their effect
is to stabilise the equilibrium. The carbon cycle, for example, is corrected by the
regulating effects of the oceans and the biological cycle, while the size of a popula-
tion is regulated by purely physical conditions (e.g. size of area available) or by the
behaviour of (groups of) organisms (e.g. competition).
The ecosystem approach was promulgated above all, however, by Howard T. and
Eugene P. Odum. E.P. Odum placed at the centre of his holistic ecology a systems
concept whose lack of clarity was thoroughly characteristic of early ecosystem
theory: on the one hand, ecosystem is a concept that describes any unit consisting
of a living component (including, for example, cells) and their environment (and
whose energy and matter transfer one is studying); on the other hand, it is also a
term that refers to a specific ecological unit (in addition to organism and popula-
tion, for example) which, as a concrete object, contains all the organisms and their
abiotic environment located in a specific area (E.P. Odum 1953, 1971; also
cf. Golley 1993). Odum postulates the necessity of a “whole-before-the-parts”
approach when studying ecosystems, because they possess emergent characteristics.
He emphasises the organismic attributes of the ecosystem and draws parallels
between succession controlled by the organic community and the development of
the individual organism; both, according to him, are oriented towards achieving
homeostasis (Odum 1969; also cf. Worster 1994; Hagen 1992, p. 128). Looking at
successive editions of their influential book “Fundamentals of Ecology” (1953,
1959, 1971) we can see how the Odum brothers increasingly built on the energetic
approach as the basis of ecology. It was Howard T. Odum in particular – likewise a
student of Hutchinson – who developed it further. He depicted the energy transfer
12 The Macy Conferences, in which figures such as N. Wiener, J. von Neumann, R. Gerard,
G. Bateson, A. Rosenblueth, M. Mead, J. von Foerster and G.E. Hutchinson participated, contrib-
uted decisively towards the dissemination of cybernetic approaches in the 1940s and 1950s far
beyond the sphere of their technical application, into areas such as the social sciences, psychology,
biology and the human and life sciences (cf. Taylor 1988; Heims 1993; Pias and Foerster 2003).
188
A. Voigt
in ecosystems using energy flow diagrams and electrical circuits (Odum 1956).
By “transforming” everything into energy he used the energetic approach as the
sole basis for researching both natural and social systems. Their energy balance
serves as well as a basis on which to evaluate them and as the starting point for
technocratic concepts of control (“ecological engineering”) – of social systems
among others (H.T. Odum 1971; for more detail, cf. Taylor 1988). Claims of this
kind to be able to explain everything and to exercise control are also found in other
systems theoretical approaches.
Ecosystem theory became the leading paradigm in ecology through various
large-scale research programmes, including studies on the distribution of radioac-
tivity, funded by the US Atomic Energy Commission, the “International Biological
Program” (IBP), undertaken in the USA from the 1960s onwards (cf. Kwa 1987;
Hagen 1992; Golley 1993) and later in Europe as well e.g. the Solling Project
(Ellenberg 1971, 1986), and the Hubbard Brook Project (Bormann and Likens
1967; Likens et al. 1977). With the emergence of the environmental movement in
the 1960s and 1970s, ecosystem research took on the task of analysing environmental
problems such as the impact of pesticides, the eutrophication of lakes etc., assessing
and combating its consequences (e.g. in the context of UNESCO’s “Man and the
Biosphere Program” (MAB). Ecosystems were to be not only studied but managed
as well. In addition, it was hoped that ecosystem theory would offer a deeper under-
standing of the effects of human action on “nature”, as well as providing, on the one
hand, an ultimately technocratic solution to the environmental crisis and, on the
other, a new, holistic human-nature relationship.
A number of factors – including the reception of different variants of systems
theory, the associated transfer of modern physical and mathematical theories into
ecology, as well as developments within the discipline – led to the flowering of a
bewildering array of ecosystem theoretical views and applications. Only a few
examples can be named here.13 Ecosystems are often described as systems charac-
terised by an open energy flow and matter cycle, and they are usually looked at on
the basis of theories of thermodynamics (e.g. Jørgensen 2000; Kay 2000).
According to this view, organisms assimilate energy in order to counter “entropic
decline”. Systems are frequently seen as being cybernetic: they can regulate themselves
physically or biologically within certain boundaries via feedback loops (e.g. Patten
1959; Patten and Odum 1981).14 The succession of ecological communities can be
described using concepts from information theory.15 The application of the hierarchy
theory in ecology (Allen and Starr 1982; O’Neill et al. 1986) means that the
ecosystem can be seen as a hierarchically organised system – neither cybernetic
models (as ultimately mechanistic models) nor the processing of ever more detailed
13 An overview of more recent developments in ecosystem theory can be found in Frontier and
Leprêtre 1998, also cf. articles in Pomeroy and Alberts 1988; Higashi and Burns 1991; Vogt et al.
1997; Pace and Groffman 1998; Jørgensen and Müller 2000; see also Chap. 27.
14 For the opposing position, cf. Engelberg and Boyarsky 1979.
15 E.g. Margalef 1958, 1968; on more modern information theoretical ecosystem approaches, see
Ulanowicz 1997; Nielsen 2000; see also Hauhs and Lange 2003.
189
15 The Rise of Systems Theory in Ecology
data are considered appropriate to complex systems. In order to understand a
phenomenon at any level of a particular hierarchy, such as that of the “ecosystem”,
it is necessary, according to this view, to look at its relationship to both higher levels
of the hierarchy (e.g. the biosphere) and lower ones (e.g. organisms). The issue of
the unpredictability of ecosystem dynamics is taken account of by referring to
catastrophe and chaos theories. A large section of the ecosystem research commu-
nity is concerned with computer modelling and simulation of ecosystems, e.g. on
the basis of fuzzy logic and artificial neural networks (cf. diverse articles in Hall
and Day 1977; Recknagel 2003). Attempts at combining population and ecosystem
ecology can be found in Jones and Lawton (1995).
The Scientization of Organicism by the Systems Concept
The different kinds of systems theories contributed to transform earlier organicist
notions of ecological units into scientific concepts, as had been the original inten-
tion of Tansley in coining the term “ecosystem”. But there is an apparent ambiguity
within the ecosystem concept.
The conception of synecological units as spatial (super)organisms (Clements
1916, 1936; Clements and Shelford 1939; Phillips 1934, 1935) implies transferring
the idea of individuality to a larger entity of organisms, to a community or organic
community. In this community individual organisms are looked at with regard to
the contribution they make towards the community’s ability to function and its
overall maintenance; its existence, characteristics and external relations are
explained by reference to the function they have for the community. In this respect,
the latter appears to be a “higher-level individual”. The ecosystem concept scien-
tizes this individual wholeness: (1) the system is “holistic” because, apart from the
“sum of its parts”, it comprises the relationships between the parts (and processes)
that, in principle, can be explained in causal terms. The ecosystem is a physical
object and is looked at as being analogous to a machine.16 The systems approach is
driven by an interest in technical knowledge to the extent that it promises the pos-
sibility of managing complex natural systems, optimising them and making them
available for use. (2) Ecosystem approaches in many cases call themselves holistic
(e.g. Odum 1953) and are indeed such, in that they take the whole of the system as
their starting point. But even so, they are reductionistic in the sense that, to a large
extent, they develop abstractions on the basis of actual objects (it was this in par-
ticular that prompted the accusation of reductionism). In the (physical) ecosystem
perspective, the main concern is with the material-energetic (and possibly informa-
tional) aspects of interactions; the actual species involved are only of interest insofar
as their specific features are relevant to the transformation of matter and energy.
Systems concepts based on set theory (e.g. Hall and Fagen 1956)17 go further still: they
16 Cf. Taylor 1988; Hagen 1992; Golley 1993.
17 “A system is a set of objects together with relationships between the objects and between their
attributes” (Hall and Fagen 1956, p. 18).
190
assert that the system’s components are not real objects but rather are defined by the
characteristics of similar classes.18 (3) With this constructivist concept of system,
“wholes” come about through a scientific operation, i.e. they are not real entities,
as in organicism.19 Ecosystems are models – theoretical constructions of the mind –
for bringing order to diverse phenomena. They are constructed according to specific
functions of the ecosystem defined by the observer (e.g. biomass production). As
such, their components are looked at from the point of view of how they contribute
towards fulfilling this function. The whole is now considered to be anything
necessary for fulfilling the function defined by the observer.
Even so, ecosystem theories may be viewed within the anti-mechanistic, organicist
tradition if, for example, it is assumed that systems have real spatial boundaries:
systems are (spatial) entities demarcated from one another by their processes; they are
subject to a succession that leads towards dynamic equilibrium, and they can be
destroyed. As such they can be depicted in terms of their relevant components, i.e. this
realistic approach assumes that the abstractions undertaken in the model correspond to
actual relationships. If the mode of functioning of an ecosystem is conceptualised in
such a way that its purpose is self-preservation, i.e. it is an end in itself (“Selbstzweck”),
then the ecosystem is being conceptualised in analogy to the organism – in other
words, a circular dependency exists between its parts; it produces, develops and main-
tains itself, and it can be destroyed. This view of ecosystems is also widely found in
nature conservation and environmental ethics.
Conclusion
In conclusion, systems theoretical views and their mathematical formulations –
therein lies both an integrating function as well as a certain set of problems – generally
allow constructivist and realist interpretations.20 A self-regulating ecosystem can be
seen as both a machine and an organism. The term “equilibrium” can be seen in
relation to something that is defined in physical-mechanistic terms, but can also be
applied to a state that is actively and intentionally maintained.21 Moreover, the idea
of the system as an organism does not stand in contradiction to the claim to capture,
manage or control it with regard to energy flow. Organicist and technocratic
perspectives may well be combined. It is between the extremes of constructivism –
realism, mechanism – organicism and reductionism – holism that the theoretical
debates within ecosystem research can be located.
A. Voigt
18 Müller 1996.
19 Cf. Tobey 1981; Jax 1998.
20 Müller 1996.
21 Cf. Weil 1999.
191
References
Allen TFH, Starr TB (1982) Hierarchy: perspectives in ecological complexity. University of
Chicago Press, Chicago
Bertalanffy L (1926) Zur Theorie der organischen ‘Gestalt’. Roux’ Archiv: 413–416
Bertalanffy L (1929) Vorschlag zweier sehr allgemeiner biologischer Gesetze. Biol. Zentralbl. 49:
83–111
Bertalanffy L (1932) Theoretische Biologie, Bd. I: Allgemeine Theorie, Physikochemie, Aufbau
und Entwicklung des Organismus. Borntraeger, Berlin
Bertalanffy L (1949) Das biologische Weltbild. Die Stellung des Lebens in Natur und Wissenschaft.
Francke, Bern
Bertalanffy L (1950) An Outline of General System Theory. Brit. J. Philos. Sci. 1:134–165
Bertalanffy L (1951) General System Theory: A New Approach to Unity of Science. Problems of
General System Theory. Human Biology 23/4:302–312
Bertalanffy L (1955) General System Theory. Main Currents in Modern Thought 11:75–83
Bertalanffy L (1968) General system theory: foundations, development applications. George
Braziller, New York
Botkin DB (1990) Discordant harmonies: a new ecology for the twenty-first century. Oxford Univ.
Pr., New York
Bormann FH & Likens GE (1967) Nutrient cycling. Science 155(3461): 424–429
Boulding KE (1941) Economic analysis. Harper & Brothers, New York
Boulding KE (1953) Toward a general theory of growth. Canadian J. o. Economics and Political
Science 19/3:326–340
Boulding KE (1956) Generals systems theory. The skeleton of science. Management Science
2:197–208
Churchman CW, Ackoff RL, Arnoff EL (1957) Introduction to operations research. Wiley,
New York
Clements FE (1916) Plant succession: an analysis of the development of vegetation. Carnegie
Institution of Washington, Washington, DC
Clements FE (1936) Nature and structure of the climax. J. of Ecology 24:252–284
Clements FE, Shelford VE (1939) Bio-ecology. Wiley, New York
Davidson M (1983) Uncommon sense: the life and thought of Ludwig von Bertalanffy, father of
general system theory. JP Tarcher, Los Angeles
Ellenberg H (ed) (1971) Integrated experimental ecology: methods and results of ecosystem
research in the German Solling Project. Springer, Berlin
Ellenberg H (ed) (1986) Ökosystemforschung. Ergebnisse des Sollingprojektes, 1966–1986.
Ulmer, Stuttgart
Engelberg J, Boyarsky LL (1979) The noncybernetic nature of ecosystems. Am Nat 114(3):
317–324
Friederichs K (1927) Grundsätzliches über die Lebenseinheiten höherer Ordnung und den ökolo-
gischen Einheitsfaktor. Naturwissenschaften 8:153–157, 182–186
Friederichs K (1934) Vom Wesen der Ökologie. – Sudhoffs Arch. Gesch. d. Medizin u.
Naturwissens 27 (3): 277–285
Friederichs K (1937) Ökologie als Wissenschaft von der Natur oder biologische Raumforschung.
Barth, Leipzig
Frontier S, Leprêtre A (1998) Développements récents en théorie des écosystèmes. Ann. Inst.
océanogr. Paris 74(1): 43–87
Gams H (1918) Prinzipienfragen der Vegetationsforschung. Ein Beitrag zur Begriffsklärung und
Methodik der Biocoenologie. Naturf. Gesellschaft Zürich. Vierteljahresschr, 63:293–493
Gerard RW (1940) Unresting Cells. Harper & Brothers, New York
Gerard RW (1953) The Organismic view of society. Chicago Behavioral Science Publications
1: 12–18
15 The Rise of Systems Theory in Ecology
192
Gleason HA (1917) The structure and development of the plant association. Bull Torrey Bot Club
44:463–481
Gleason HA (1926) The individualistic concept of the plant association. Bull Torrey Bot Club
53:7–26
Golley FB (1993) A history of the ecosystem concept in ecology: more than the sum of the parts.
Yale University Press, New Haven/London
Hagen JB (1992) An entangled bank: the origins of ecosystems. Chapman & Hall, New York
Hall CAS, Day J (eds) (1977) Ecosystem modeling in theory and practice. Wiley, New York
Hall AD, Fagen RE (1956) Definition of System. General System, 118–28
Hammond D (2003) The science of synthesis: exploring the social implications of General
Systems Theory. Univ. Pr. of Col., Colorado
Hauhs M, Lange H (2003) Informationstheorie und Ökosysteme. Handbuch der
Umweltwissenschaften. Ecomed, München: 1–22
Heims SJ (1993) Constructing a social science for postwar America: the cybernetics group,
1946 – 1953. MIT Press, Cambridge
Higashi M, Burns TP (eds) (1991) Theoretical studies of ecosystems. Cambridge University Press,
Cambridge
Hutchinson GE (1948) Circular causal systems in ecology. Annals of the New York Academy of
Sciences 50:221–246
Jax K (1998) Holocoen and ecosystem: on the origin and historical consequences of two concepts.
J. Hist. Biology, 31:113–142
Jax K (2002) Die Einheiten der Ökologie. Analyse, Methodenentwicklung und Anwendung in
Ökologie und Naturschutz. Lang, Frankfurt/M
Jones CG, Lawton JH (1995) Linking species and ecosystems. Chapman & Hall, New York
Jørgensen SE (2000) A general outline of thermodynamic approaches to ecosystem theory. In:
Jørgensen S, Müller F (eds) Handbook of ecosystem theories and management. Lewis,
London/New York/Washington, DC
Jørgensen SE, Müller F (2000) Handbook of ecosystem theories and management. Lewis,
London/New York/Washington, DC
Kay JJ (2000) Ecosystems as self-organising holarchic open systems: narratives and the second
law of thermodynamics. In: Jørgensen S, Müller F (eds) Handbook of ecosystem theories and
management. Lewis, London/New York/Washington, DC
Köhler W (1920) Die physischen Gestalten in Ruhe und im stationären Zustand: eine naturphil-
osophische Untersuchung. Vieweg, Braunschweig
Kwa C (1987) Representations of nature mediating between ecology and science policy: the case
of the International Biological Programme. Social Studies of Science 17, 3, 413–442
Lamotte M, Bourliere F (1978) Problemes d’ écologie, structure et fonc-tionnement des écosys-
tèmes terrestres. Masson, Paris
Lotka, AJ (1925) The elements of physical biology. Williams & Wilkins, Baltimore
Lindeman RL (1942) The trophic-dynamic aspect of ecology. Ecology 23:339–418
Likens GE, Bormann FH, Pierce RS, Eaton JS, Johnson NM (1977) Biogeochemistry of a forested
ecosystem. Springer, New York
Lilienfeld R (1978) The rise of systems theory. Wiley, New York
Margalef R (1958) Information theory in ecology. YearB Soc Gen Syst Res 3:36–71
Margalef R (1968) Perspectives in ecological theory. University of Chicago Press, Chicago,
pp 1–25
Maturana HR & Varela FJ (1987) Der Baum der Erkenntnis: die biologischen Wurzeln des men-
schlichen Erkennens. Scherz Verlag, Bern
McIntosh RP (1985) The background of ecology: concept and theory. Cambridge University
Press, Cambridge
McIntosh RP (1995) H. A. Gleason’s ‘Individualistic concept’ and theory of animal communities:
a continuing controversy. - Biol. Rev., 70:317–357
Müller K (1996) Allgemeine Systemtheorie. Studien zur Sozialwissenschaft 164. Opladen
A. Voigt
193
Neumann J, Morgenstern O (1944) Theory of games and economic behavior. Princeton Univ.
Press, Princeton, NJ
Nielsen SN (2000) Ecosystems as information systems. In: Jørgensen S, Müller F (eds) Handbook
of ecosystem theories and management. Lewis, London/New York/Washington, DC
Odum E (1953, 1959, 1971) Fundamentals of ecology. Saunders, Philadelphia
Odum HT (1956) Primary production in flowing waters. Limnology and Oceanography
1:102–117
Odum EP (1969) The strategy of ecosystem development: an understanding of ecological succes-
sion provides a basis for resolving man’s conflict with nature. Science 164:262–270
Odum HT (1971) Environment, power and society. Wiley, London
O’Neill RV, DeAngelis DL, Waide JB, Allen TFH (1986): A hierarchical concept of ecosystems.
Princeton Univ. Pr., Princeton, NJ
Parsons T (1937) The structure of social action. McGraw-Hill, New York
Pace ML, Groffman PM (eds) (1998) Successes, limitations, and frontiersn in ecosystem science.
Springer, New York
Patten BC (1959) An introduction to the cybernetics of the ecosystem: the trophic dynamic aspect.
Ecology 40:221–231
Patten BC, Odum EP (1981) The cybernetic nature of ecosystems. Am Nat 118:886–895
Peus F (1954) Auflösung der Begriffe “Biotop” und “Biozönose”. Deutsche Entomologische
Zeitschrift N F 1:271–308
Phillips J (1934,1935) Succession, development, the climax, and the complex organism: an analy-
sis of concepts. Part 1–3. J Ecol 22:554–571, 23: 210–246¸ 3: 488–508
Pias C & Foerster H (eds) (2003) Cybernetics: the Macy-Conferences 1946–1953. Diaphanes,
Zürich
Pomeroy LR, Alberts JJ (eds) (1988) Concepts of ecosystem ecology. Springer New York
Prigogine I (1955) Introduction to thermodynamics of irreversible processes. Thomas,
Springfield
Ramensky LG (1926) Die Gesetzmäßigkeiten im Aufbau der Pflanzendecke. Botanisches
Centralblatt N F 7:453–455
Rapoport A (1947) Mathematical theory of motivation of interactions of two individuals. Bulletin
of Mathematical Biophysics 9,1:17–27
Rapoport A (1950) Science and the goals of man: a study in semantic orientation. Harper,
New York
Recknagel F (ed) (2003) Ecological informatics: understandig ecology by biologically-inspired
computation. Springer, Berlin
Shannon CE, Weaver W (1949) The mathematical theory of communication. University of Illinois
Press, Urbana, Illinois
Schwarz AE (1996) Aus Gestalten werden Systeme: Frühe Systemtheorie in der Biologie. In: Mathes
K, Breckling B, Eckschmitt K (eds) Systemtheorie in der Ökologie. Landsberg, pp 35–45
Tansley AG (1935) The Use and abuse of vegetational concepts and terms. Ecology 16(3):
284–307
Taylor P (1988) Technocratic optimism, H.T. Odum, and the partial transformation of ecological
metaphor after World War II. – J. Hist. Biol., 21(2):213–244
Thienemann A, Kieffer JJ (1916) Schwedische chironomiden. Arch. hydrobiol. 2(Suppl):489
Tobey RC (1981) Saving the prairies. University of Carlifonia, Berkeley
Trepl L (1987) Geschichte der Ökologie. Vom 17. Jahrhundert bis zur Gegenwart. Athenäum,
Frankfurt a. M.
Ulanowicz RE (1997) Ecology, the ascendent perspective. Columbia University Press,
New York
Vogt KA, Gordon JC, Wargo JP, Vogt DJ, Asbjorsen H, Palmiotto PA, Clark HJ, O’Hara JL,
William S-K, Toral P-W, Larson B, Tortoriello D, Perez J, Marsh A, Corbett M, Kaneda K,
Meyerson F, Smith D (1997) Ecosystems: balancing science with management. Springer,
New York
15 The Rise of Systems Theory in Ecology
194
Voigt A (2001) Ludwig von Bertalanffy: Die Verwissenschaftlichung des Holismus in der
Systemtheorie. Verhandlungen zur Geschichte und Theorie der Biologie 7:33–47
Volterra V (1926) Variazioni e fluttuazioni del numero d’individui in specie animali conviventi.
Mem. Accad. Lincei series 6, 2(36):31–113
Weil A (1999) Über den Begriff des Gleichgewichts in der Ökologie - ein Typisierungsvorschlag.
Unversitätsverlag, TU Berlin, Berlin
Wiener N (1948) Cybernetics or control and communication in the animal and the machine.
Wiley, New York
Worster D (1994) Nature’s economy: a history of ecological ideas. Camb. Univ. Pr., Cambridge
A. Voigt
195
Chapter 16
Ecology and the Environmental Movement
Andrew Jamison
Introduction
With the emergence of the environmental movement in the 1960s, the science of
ecology was transformed from being a relatively minor sub-field of biology into an
object of political engagement and public interest. For a brief historical moment,
ecology became more than a mere science; it became a component part of what
I have previously characterized as an emerging ecological culture (Jamison 2001).
Even though many of the political struggles that brought it into being have faded into
the past, the environmental movements of the 1960s and 1970s continue to influence
scientific ideas and personal values, as well as broader socio-political discourses.
Ecology became a kind of “super-science” that was considered to have a crucial
role to play in the newfound mission, or political project, of environmental protec-
tion. From various points along the political spectrum, it was felt that the science
of ecology could provide concepts, theories, and methods that could help guide
society into a more sustainable, environmentally-friendly direction. These develop-
ments were initiated in the United States but spread to Europe in the course of the
1970s as environmental movements took shape in many countries. The influence of
a politicized ecology was particularly strong in the Scandinavian countries, where
there were indigenous traditions of both ecological science and environmental politics
dating back to the eighteenth century.
Opinions varied, however, in regard to just what the science of ecology had to
offer to the broader politics of the environment. According to the distinction made
by the Norwegian philosopher Arne Naess in 1972, there was both a “shallow” and
a “deep” version of ecology that could be found in what was starting to be consid-
ered an environmental movement. Where the shallow ecologists went to the science
primarily in search of operational concepts and administrative tools with which to
carry out their political struggles, the deep ecologists took their point of departure in
A. Jamison (*)
Department of Development and Planning, Institut for Samfundsudvikling og Planlægning,
Aalborg University, Fibigerstræde 13, 9220 Aalborg, Denmark
e-mail: andy@plan.aau.dk
A. Schwarz and K. Jax (eds.), Ecology Revisited: Reflecting on Concepts, Advancing Science,
DOI 10.1007/978-90-481-9744-6_16, © Springer Science+Business Media B.V. 2011
196
A. Jamison
the science for the creation of a new world-view, or belief system, an ecological
philosophy. In a similar vein, the American activist Murray Bookchin distinguished
between what he termed “environmentalists” and “ecologists” within the emerging
movement (e.g. Bookchin 1982). Bookchin’s argument was that the environmentalists
reacted to particular cases of environmental destruction, while the ecologists reacted
to the underlying social and political conditions behind the particular cases. Others
referred to a tension in the emerging environmental consciousness between ecocen-
trists and anthropocentrists, which was due, in large measure, to different meanings
that were given to the science of ecology in its relation to environmental politics.
Wherever environmental movements developed, the science of ecology came to
play a significant role in the collective identity, or cognitive praxis, of the activists
and their activities. Ecology, in various ways, took on an ideological function, rather
than, or in addition to, its more traditional scientific role in society. The “use” of
ecology for political purposes proved to be problematic, however, and, in the course
of the 1980s, the science of ecology and the environmental movement more or less
parted company. The discourse of sustainable development tended to replace ecology
as an overarching ideology, or political doctrine, for environmental activists and
green party politicians, and most ecologists tended to disavow, or at least, disassociate
themselves from the explicit political meanings that had been attributed to their
science. But the links that were established in the 1970s have continued to affect
both the science of ecology, as well as the broader politics of the environment.
Indeed, in the recent challenges to the scientific understanding of climate change
from self-declared skeptics such as the Danish political scientist Bjørn Lomborg,
the views of environmental scientists and environmental activists are conflated with
one another. For skeptics such as Lomborg, the statements of ecologists and other
environmental scientists are not to be believed, at least in part because of their asso-
ciation with environmental organizations, such as Greenpeace (Lomborg 2001).
The aim of this article is to discuss the interactions between ecology and the
environmental movement with particular reference to the way that ecology came to
be a kind of “super-science” in the 1960s and 1970s.
The Traditions of Ecology
In an influential account, first written in the 1970s, Donald Worster pointed to two
main streams of thought that had come together in the environmental movement, two
opposing attitudes to nature that had led, through the centuries, to two different kinds
of ecology, or ecological traditions (Worster 1979). He traced an “imperialist” tradi-
tion back to the writings of Francis Bacon in the early seventeenth century and his
ideas about the human domination of nature. Carl Linneaus and Georges Buffon in
the eighteenth century helped give this imperialist ecology a more systematic and
scientific form. Nature was conceptualized in mechanical and instrumental terms,
which helped make possible the effective utilization of nature for human exploitation.
In the course of industrialization, this highly utilitarian view of non-human reality
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16 Ecology and the Environmental Movement
became the dominant discourse, or philosophy, of nature, especially in the natural
sciences, as they took on a more professional and disciplined organizational identity.
The Linnean system provided methods that were used both for scientific research
and for more technical kinds of work. The various plants and animals were given
names, which made it easier to understand and analyze their functions and inter-
relations, and they were also given structural characteristics, which proved useful for
conducting practical experiments, such as breeding new plant varieties or exploiting
new natural resources. The imperialist tradition represented the experimental, sys-
temic approach to knowledge-making, which entered the science of ecology, when
it was given a more formalized identity as a sub-field of biology around the turn of
the century in both the United States and several European countries.
Opposed to the imperialists were the nature-lovers, to whom Worster gave the
label “arcadian” in order to associate their version of ecology to the classical ideal
of harmony between nature and society that had been depicted by Roman poets in
the Greek region of Arcady. According to Worster, the back-to-nature folks began
to articulate their counter-program at the dawning of the industrial era, as part of
the Romantic movement. The arcadians shared many of the modernizing, scientific
ambitions of the imperialists, but they came to develop a different way of investi-
gating and understanding nature. Tracing arcadians back to the English pastor and
writer Gilbert White, and especially to his work, “The Natural History of Selborne”,
originally published in 1789, Worster delineated a stream of experiential, or partici-
patory, ecology that was perhaps most influentially developed further by Henry
David Thoreau in the nineteenth century. In Germany and the Scandinavian coun-
tries, a related tradition of “Naturphilosophie” (or philosophy of nature) emerged
in the late eighteenth century among academics and artists; this romantic tradition
had an influence in both geology and geography, biology and chemistry, and even
in physics, where the search for an underlying “spirit” in nature led the Danish
scientist Hans Christian Ørsted to discover the connection between electricity and
magnetism in 1820.
Worster’s argument was that the two ecological traditions had both contributed
to Charles Darwin’s theory of natural evolution, but that they had subsequently
given rise, in the course of the twentieth century, to two different ways of thinking
about ecology and conducting ecological research. The one was systemic, while the
other was individual in focus, and they fostered a large-scale, ecosystems oriented
ecology, on the one hand, and a smaller-scale, population-oriented ecology on the
other, the one taking its point of departure in the systemic relations that exist among
species, and the other taking its point of departure in the dynamic relations of one
species to its environment. The two traditions drew on different attitudes, or
conceptions of nature, as well as different methodological and theoretical assumptions
about how to investigate, or interrogate nature.
Worster’s division into an imperialist and arcadian ecology captures a funda-
mental contradiction in the history of ecology. But it tends to disregard a third
important source of inspiration for the environmental movement, as it developed in
the 1960s and 1970s, namely the various “human ecologies” that had emerged in the
nineteenth and early twentieth centuries, both in Europe and in the United States.
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A. Jamison
In part motivated by the journeys of exploration among biologists and geographers
to South America and in the North American frontier, in part an outgrowth of infra-
structural projects and urban planning and, in part, a sub-field of public medicine
and public health, these human ecologies entered into the new social sciences of
sociology and anthropology, of economics and political science, and of geography
and planning as they grew into important fields in the course of the twentieth
century.
To make the story somewhat more complete, it can therefore be useful to add a
third tradition to Worster’s two, and to distinguish three ideal-typical ecological
traditions that have been mobilized in the making of the environmental movement.
Each tradition – the imperialist, the arcadian, and the human – has its own charac-
teristic conception of nature and its own preferred methods of investigation, as well
as its own distinct version of an appropriate ecological practice or politics )
Table 16.1).
The Mobilization of Traditions
It would be these three traditions and the various sub-sets thereof that came to be
mobilized in the 1960s in the making of a new social movement. On the one hand,
the imperialist tradition was reinvented, among other places, in the cybernetic language
of ecosystems ecology and energy systems analysis (see Chap. 15). Systems ecology,
as developed by Eugene and Howard Odum, became extremely influential among
natural scientists, particularly during the International Biological Program, and, as
a new approach to ecology, it would play a major role in the emergence of an envi-
ronmental consciousness in the 1960s (Worster 1979, pp. 291ff.).
The Odums also illustrate the importance of established scientists in the articulation
of the new environmentalism’s collective identity, or cognitive praxis (Cramer et al.
1987). The environmental movement involved, at the outset, a kind of popularization
of science, as well as a translation of concepts and terminology that had been
Table 16.1 Ecological traditions
Imperialist Arcadian Human
Formative
influences
Francis Bacon
Carolus Linnaeus
Gilbert White
Henry D Thoreau
George Marsh
Lewis Mumford
Key mobilizers Odum brothers
Gro H Brundtland
Rachel Carson
Arne Næss
Paul Ehrlich
Barry Commoner
Type of sciencing Systemic models
experimentation
Natural history thick
description
Mapping surveying
Relation to nature Exploitation management Participation harmony Planning
co-construction
Conception of nature Ecosystem resource base Community locality Region landscape
Ideologies Anthropocentric/
modernism
Ecocentric/deep
ecology
Pragmatic/
postmodern
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16 Ecology and the Environmental Movement
developed in understanding non-human nature to human societies. Eugene and
Howard Odum’s popular writings provided a scientific legitimacy and authority to
the new movement, as well as a powerful terminology and conceptual framework,
while the movement helped provide ecological scientists with new opportunities for
research, and a new political mission: ecologizing society (Söderqvist 1986).
In the World Wildlife Fund, founded in 1961, and then more scientifically in
the International Biological Program, that was established in the mid-1960s, the
“imperialist” tradition took on a more modern or contemporary manifestation. It
became more explicitly international, as scientists and other conservationists came
to take part in transnational research and development networks, particularly within
the IBP projects (Kwa 1989). The imperialist tradition was also brought up to date
technologically with the new cybernetic and computer-based approaches to research
that were developed by the new breed of ecosystem ecologists, led by Eugene and
Howard Odum. Particularly in the energy-flow schematizations of Howard Odum,
the mathematical modeling of nature and society was presented in an ambitious and
sophisticated manner that was to have a major importance on the emerging environ-
mental consciousness.
It would be the biologist turned science writer Rachel Carson, whose eloquent
writings would do most to give the arcadian tradition a contemporary resonance.
“Over increasingly large areas of the United States”, she wrote, “spring now comes
unheralded by the return of the birds, and the early mornings are strangely silent
where once they were filled with the beauty of bird song” (Carson 1962, p. 97). Her
book “Silent Spring” served to awaken the industrial world from its postwar slumbers,
and she was soon followed by other writers who, with their scientific pondus and
more sober tone, had a somewhat different impact on the public consciousness.
After writing two best-selling nature books in the 1950s, Carson had grown
concerned about the impact that the new chemical insecticides were having on the
forests and on the animals that she loved so much. Her 4-year investigation of the
environmental consequences of one of those pest-killers resulted in a new form of
political broadside, a book of scientific poetry that brought the arcadian science
of Thoreau, whose writings Carson admired so much, into the twentieth century. In
any case, “Silent Spring” was to have a major influence on the cognitive praxis of the
emerging environmental movement (Jamison and Eyerman 1994, pp. 92ff.). But it
would also inspire a new generation of arcadian ecologists to reframe their message
and challenge the more “technocratic” approaches of the systems ecologists. To a large
extent, the historical dichotomy between the imperialists and the arcadians would be
replayed in the tensions over direction and orientation in the fledgling environmental
movement organizations that developed in the late 1960s and early 1970s.
The mobilization of human ecology came from many different directions. Some,
like Murray Bookchin, who had been a labor activist in the 1930s, brought a socialist
sensibility into the environmental movement. His book from 1963, “Our Synthetic
Environment”, was one of the first to present the wide range of new environmental
problems – occupational health, chemical pollution, household risks, waste disposal –
that were to gain increasing public attention in the years to come. Others, like the
biochemist Barry Commoner, gave the environmental movement a more technical
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A. Jamison
emphasis; Commoner depicted, in his first book, “Science and Survival” (1966), the
subservient role that science was playing in society and production, and suggested a
number of public service, or critical, activities for scientists to play in the emerging
movement. The biologist Paul Ehrlich resurrected the Malthusian message of
population pressures and resource limitations in his book, “The Population Bomb”
(1968), and the different perspectives of Commoner and Ehrlich would subse-
quently combine in new activist organizations and environmental studies departments.
Still others, like Lewis Mumford, would provide historical and philosophical
perspectives to help understand the new environmental problems. As such, the
human ecology tradition was also reinvented, or mobilized, in the 1960s.
Already then, however, the seeds were sewn for the differentiations that have
followed. The mobilization of traditions did not lead to one coherent movement, but
rather to different kinds of hybrid identities which, for the sake of simplicity, we
can think of as practical, cultural, and political. The practical, or technical environ-
mentalists, have reinvented the human ecological tradition with infusions of
advanced technology, but also with influences from the other ecological traditions.
And while the cultural and the political environmentalists have both identified most
strongly with one of the earlier traditions, they too have combined the older idea
and perspectives with new ingredients. For the culturalists, the hippie influence,
and the more general critique of technocratic society and its “one-dimensional”
thought, have been of fundamental importance; while, for the political environmen-
talists, the “globalizing” tendencies of capitalism, and the development of telecom-
munications and media technologies have been extremely significant factors in
their professionalization.
The Age of Ecology
By the end of the 1960s, ecology had inspired both the emergence of new activist
groups, such as Friends of the Earth, as well as a process of policy reform and
institution building. In the early 1970s, most of the industrialized countries estab-
lished new state agencies to deal with environmental protection, and environmental
research and technological development were organized in new locations in both
the private and public sectors – often in the name of ecology. Many national parliaments
enacted more comprehensive environmental legislation and, at the United Nations
Conference on the Human Environment in Stockholm in 1972, protecting the environ-
ment was recognized as a new area of international concern.
The manned landing on the moon in 1969 had provided the symbol for the
conference, the blue planet viewed from space: small, fragile, and strikingly beautiful
in its shape and color. A biologist, René Dubos, and an economist, Barbara Ward,
collaborated on the book that would set the agenda for the conference. “Only One
Earth” (1972), their book was called, and in it they made the case for a new kind of
environmentalism, combining efficient management of resources with empathetic
understanding: “Now that mankind is in the process of completing the colonization
of the planet”, they wrote, “learning to manage it intelligently is an urgent imperative.
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16 Ecology and the Environmental Movement
Man must accept responsibility for the stewardship of the earth” (Ward and Dubos
1972, p. 25). They noted, in conclusion, that the reforms and policy proposals that
they suggested in their book would not come easily: “The planet is not yet a centre
of rational loyalty for all mankind. But possibly it is precisely this shift of loyalty
that a profound and deepening sense of our shared and inter-dependent biosphere
can stir to life in us” (Ward and Dubos 1972, p. 298).
Over the next few years, ecology would come to be drawn upon by almost all sides
in the new environmental politics. There was also a range of “grass-roots” engineer-
ing initiatives that emerged in the fledgling environmental movement, a kind of eco-
logical technology. In the United States, a group of self-proclaimed “new alchemists”
moved from the university out to the country to experiment with ecological agricul-
ture and renewable energy (Todd 1977). In many European countries, but perhaps
especially in Britain, Denmark and the Netherlands, a number of research centers and
projects in alternative or ecological technology were established. Also in Switzerland
and in Germany, the so-called anthroposophic movement of Rudolf Steiner, that had
been started in the 1920s with its biological-dynamic agri- and horticulture was rein-
vigorated, as were the organic-biological methods invented by the Swiss Hans Müller
in the 1950s. At some of the “hippie” communes and production collectives that
developed at the time, there was often an interest in energy and agriculture, and there
were also, among architects and planners, attempts to develop more environmentally-
friendly, or ecological approaches and techniques (Dickson 1974).
An interest in ecological science and technology developed as an integral part of
the environmental movement in several countries, as ecological research was given
greatly increased funding, and programs in environmental education were estab-
lished at many universities. In retrospect, we can see that the environmental move-
ment opened a public space for experimentation with a collective mode of science
and engineering – or what Ivan Illich called at the time “tools for conviviality” (Illich
1973) – in relation to energy, agriculture, housing, and transportation. The particular
technical interests have diffused widely into society – for are we not all a bit more
“ecological” in the ways we garden, and decorate our homes, and move ourselves
around? – while the collective creativity has largely dissipated. While new scientific
institutions eventually emerged out of the movement, most of the attempts that were
made in the early 1970s to “use” ecology directly for purposes of environmental deci-
sion making proved unsuccessful. But even more significant in the long run was the
coming of the oil crisis and the central role that the struggle over nuclear energy
came to play in the second half of the 1970s in both Europe and North America.
New kinds of disciplines, or sub-disciplines developed in many countries.
Energy systems analysis became a recognized field for investigating the costs and
benefits of different choices of energy supply and distribution. Human ecology took
on the form of a recognized academic field, and developed its own theories, based
on concepts of entropy and energy flow. Within ecology itself, a kind of bifurcation
took place between ecosystem ecologists, on the one hand, who were often drawn
into larger, multidisciplinary projects, and population, or evolutionary ecologists, on
the other, who focused their attention on particular species or ecological communities.
There was a still further specialization, due to the range of approaches that emerged
in the established disciplines to take on the new environmental and energy issues.
202
The Politicization of the Movement
It was the first oil crisis, of 1973–74, that led to a major shift in environmental
consciousness, as energy issues moved to the top of many national political agen-
das, especially in relation to nuclear energy. In many countries, the late 1970s were
a period of intense political debate and social movement activity, as the pros and
cons of nuclear energy, or “hard energy paths” in general, were contested (Lovins
1977). In certain countries, like Denmark, renewable energy experimentation
became a social movement of its own, and led to new industries and government
programs. Seen in retrospect, an important result of the energy debates of the 1970s
was a professionalization of environmental concern and an incorporation by the
established political structures of what had originally been a somewhat delimited
political issue. As a result, there was both a specialization and institutionalization
of knowledge production.
What, in retrospect, is most characteristic of the 1970s is the breadth, but also
the unity and coherence, of the environmental movement. As a popular front, or
campaign, against nuclear energy, the different traditions of ecology were com-
bined into an integrative cognitive praxis, with a visionary ecological philosophy,
or world-view guiding a range of practical experiments with alternative technology
in settings that were largely autonomous, outside of the formalized rule systems
and organizational frameworks of the larger society. In informal local groups, and
movement-based workshops and study circles, technical projects and educational
activities were conducted with participation of both “experts” and amateurs. The
key point is that the movement, for a brief time, could provide an organized learning
experience, in which theory and practice were combined in pursuit of a common,
collective struggle. These settings would be difficult to maintain for very long,
since they were, in many ways, too unstable for any kind of permanent institution-
alization, and when the issues that inspired the movement were resolved, and taken
off the political agenda, the different component parts split apart and fragmented
(Cramer et al. 1987). The unity that had been achieved in struggle could simply not
be sustained.
The challenge to the coherence of the movement was also, to a large extent, a
result of the broadening and diversification of environmentalism in the late 1970s.
While most activists in Europe were concerned with nuclear energy, which became
an issue of major political importance in several countries, other issues were impor-
tant and inspired new forms of activism in other parts of the world. In the United
States, the discovery of toxic wastes buried under the neighbourhood of Love Canal
in Buffalo, New York, inspired a new kind of locally-based, working class opposi-
tion to environmental pollution (Szasz 1994). It was also in the United States that
the new techniques of genetic engineering were critically reviewed by activists for
their risks and dangers to the communities in which the laboratory experiments
were carried out. In opposing genetic engineering, environmentalists, such as
Jeremy Rifkin, pointed to a new kind of futuristic challenge that made many of the
actual environmental problems pale in significance. “Two futures beckon us”,
A. Jamison
203
References
Bookchin M (1982) The ecology of freedom. Cheshire Books, Palo Alto
Carson R (1962) Silent spring. Houghton Mifflin, Boston
Cramer J, Eyerman R, Jamison A (1987) The knowledge interests of the environmental movement
and its potential or influencing the development of science. In: Blume S, Bunders J, Leydesdorff
L, Whitley RP (eds) The social direction of the public sciences. Reidel, Dordrecht
Dickson D (1974) Alternative technology and the politics of technical change. Fontana, Glasgow
Rifkin wrote, “[w]e can choose to engineer the life of the planet, creating a second
nature in our image, or we can choose to participate with the rest of the living
kingdom. Two futures, two choices. An engineering approach and an ecological
approach” (Rifkin 1983, p. 252).
These new forms of environmentalism were difficult to contain within one unified
movement; rather, in the intellectual and broader political traditions that they
tapped into, and in the alliances that they made, sometimes with quite conservative
and religiously fundamental groups, they were often articulating interests and strategies
that were diametrically opposed to the positions of “modernist” anti-nuclear activists,
as well as many of the professional environmentalists in the think tanks and the
“mainstream organizations”. As such, while the environmental movement was
growing and expanding and diversifying, the seeds were sewn for a more explicit
process of differentiation in the 1980s.
The tension between those who see the environmental crisis as fundamental and
those who see it as merely one of many challenges facing modern society has been a
defining feature of the environmental movement ever since, and it has exerted a strong
influence on the way in which ecology has been used, or appropriated. The “funda-
mentalists” or deep ecologists tend to see ecology as an overarching philosophy or
cosmology, while the pragmatists or realists often see ecology as a somewhat more
limited scientific toolbox, providing methods and concepts for environmental managers
and other scientists. Perhaps the most appropriate approach for environmentalists to
follow in the future is to try to forge a new kind of hybrid identity that combines the
passion, engagement and holistic thinking of the deep ecologists with the practical
skills and professional rigor of the environmental managers.
Indeed, it might be suggested that it is in the hybridization of knowledge-making
activities that the environmental movement, or environmental activism continues to
have an impact on the development of ecology, and, more generally on the making of
“green knowledge” (Jamison 2001). In recent years, in the quest for sustainable
development, there have emerged a number of new cognitive combinations or trans-
disciplinary forms of knowledge production that make connections between the natural,
social, and human sciences (Jamison 2005). As such, ecology has once again come
to play a broader role in society, both in environmental politics and management (in
such hybrid practices as industrial ecology and ecological economics), as well as
within the larger culture (in relation to such hybrid “discourses” as ecological citizen-
ship and ecological literacy).
16 Ecology and the Environmental Movement
204
Illich I (1973) Tools for conviviality. Harper & Row, New York
Jamison A (2001) The making of green knowledge. Environmental politics and cultural
transformation. Cambridge University Press, Cambridge
Jamison A (2005) Hybrid identities in the european quest for sustainable development. In: Paehlke
R, Torgerson D (eds) Managing leviathan. Broadview Press, Peterborough
Jamison A, Eyerman R (1994) Seeds of the sixties. University of California Press, Berkeley
Kwa Chunglin (1989) Mimicking nature. The Development of Systems Ecology in the United
States, 1950–1975. Dissertation, University of Amsterdam, Amsterdam
Lomborg B (2001) The skeptical environmentalist. Cambridge University Press, Cambridge
Lovins A (1977) Soft energy paths. Penguin, Harmondswoth
Rifkin J (1983) Algeny. Penguin, Harmondsworth
Szasz A (1994) Ecopopulism: toxic waste and the movement for environmental justice. University
of Minnesota Press, Minneapolis
Söderqvist T (1986) The ecologists. From merry naturalists to saviours of the nation. Almqvist
and Wiksell, Stockholm
Todd NJ (ed) (1977) The book of the new alchemists. E.P. Dutton, New York
Ward B, Dubos R (1972) Only one earth. The care and maintenance of a small planet. Andre
Deutsch, London
Worster D (1979) Nature’s economy. The roots of ecology. Anchor Books, Garden City
A. Jamison
205
Chapter 17
Ecology and Biodiversity at the Beginning
of the Twenty-first Century: Towards a New
Paradigm?
Patrick Blandin
Introduction
Since the 1960s, Ecology is a matter of crucial concern, with people becoming
increasingly aware of the increasing environmental crisis. In reality, as a scientific
discipline, Ecology was first officially addressed when the International Union for
the Protection of Nature (IUPN, now IUCN, with “C” for “Conservation”) was
created at the end of 1948: an agenda was drawn up, including the following topic:
“the international cooperation for scientific research in the field of the Protection of
Nature, especially concerning œcological research in the various fields of exact and
natural sciences” (UIPN 1948, p. 15). The first concrete IUPN action was the orga-
nization, with UNESCO, of a technical conference, which took place at Lake
Success (USA), in August, 1949. The Ecology Section of the conference was intro-
duced by a French biologist, Georges Petit, who emphasized the fact that the
relationships between the Protection of Nature and Ecology had been widely
neglected, the Protection of Nature having been “considered for a long time only as
the results of aesthetic or moral preoccupations” (Petit 1950, p. 304).
It is important to grasp the conceptual situation at this very moment when the
conservationist movement met Ecology. The IUPN first General Secretary’s view-
point (Harroy 1949, p. 10) illustrates conservationists’ expectations: “In order to
efficiently protect the natural associations which are useful, Man must have them
carefully studied beforehand. But to study these associations in the best conditions,
I would say “in the state of a pure body”, he must have protected them before, that
is, in appropriate and sufficiently vast areas, to have them shielded from disturbing
human influences which mask and distort the fundamental processes that the
researcher attempts to observe and to order into laws”. This is symptomatic of an
ideology which considers man as an external factor, whose actions disturb the natu-
ral equilibrium – the so called “Balance of Nature”: – an ideology which is the
descendant of Bacon’s and Descartes’s dualist philosophies. Rapidly, this ideology
P. Blandin (*)
Muséum National d’Histoire Naturelle, Départment Hommes-Natures-Sociétés, 57, rue Cuvier,
75005 Paris, France
e-mail: patrick.blandin@yahoo.fr
A. Schwarz and K. Jax (eds.), Ecology Revisited: Reflecting on Concepts, Advancing Science,
DOI 10.1007/978-90-481-9744-6_17, © Springer Science+Business Media B.V. 2011
206
P. Blandin
fit with the developing systemic and cybernetic approach of Ecology, which bloomed
under the umbrella of the Odumian Ecosystem Paradigm (Bergandi 1995).
At the turn of the twentieth century, the environmental crisis – of which the
Biodiversity accelerated loss and the climatic change are emblems – radically questions
the comfortable dualist occidental ideology: new ways of living with Nature are to
be invented. In this context, the Biodiversity challenge calls for the renewal of
Ecology, which is in search of a paradigm for the ongoing twenty-first century. In
this paper, I shall briefly analyse this evolution of Ecology, focussing on the issue
of biological diversity.
The Balance of Nature Ideology and the Ecosystem Stability
At the Lake Success IUPN Conference, the term “ecosystem” (Tansley 1935) was
not used: the prevailing concern was about the consequences of different human
activities on the “Balance of Nature”; the ecosystem concept came into common
use later, after the publication of Odum’s “Fundamentals of Ecology” (1953). In
fact, Ecosystem Ecology developed independently of conservation issues, but
shared with the conservation world the Balance of Nature Ideology, which claims
that equilibrium is the “normal” state of Nature.
Rapidly, in the USA, thanks to the availability of the first digital computers, the
use of ecosystem analysis and cybernetics models became the core of ecological
research for decades (Golley 1991). For many ecologists, the ecosystem being con-
ceived as an organized, “cybernetic” entity, its biotic community could not be con-
sidered as a random assemblage of species. The community would present a
“structure”, determined by interactions between the coexisting species – the number
of which being probably ecologically regulated, depending on available resources to
be shared by competing species –, and resulting in global properties at the ecosys-
tem level. The Odumian Ecosystem Paradigm implies that the interactions between
the components of any natural ecosystem, playing under the control of external and
internal constraints, result in the stability of the mature “climax” ecosystem. Little
by little, the intuitive idea of some relationships between species diversity, structural
and functional complexity, and ecosystem stability naturally emerged. Therefore,
during the 1960s and the 1970s, much research focussed on the structure of ecologi-
cal communities, and many papers dealt with the problem of the relationships
between species diversity and ecosystem characteristics and stability (for example:
Leigh 1965; Paine 1966; Margalef 1969; Loucks 1970 or Smith 1972).
In 1973, in his Presidential address to the British Ecological Society, Amian
Macfadyen emphasized the fact that the question of species diversity – ecosystem
stability relationships being controversial, Ecology was still in an early stage and
lacked accepted paradigms (Macfadyen 1975). The question was reviewed by
Daniel Goodman (1975), who observed that “It would seem, at first sight, to be
plausible that the balance of nature is more readily balanced when there are more
interacting species present. This functional relationship may be conceived in terms
of spare parts, more links to take up the slack, or, nowadays, more opportunity for
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17 Ecology and Biodiversity at the Beginning of the 21st Century
feedback loops.” (p. 238). But Goodman concluded that “the expectations of the
diversity-stability hypothesis are borne out neither by experiments, by observation,
nor by models” (p. 261). Actually, at the end of the 1970s the interest in species
diversity as a global characteristic of communities was decreasing, perhaps because
it was actually impossible to develop “popperian” research on the supposed
ecological functions of species diversity.
Biodiversity Appears on the Scene
Diversity began a new life at the USA National Forum on BioDiversity, held in
1986, which had an immediate impact on the public and the media. Edward
O. Wilson, in his Editor’s foreword of the resulting book, entitled “BioDiversity”,
(1988a, p. v), pinpointed two “more or less independent developments” to explain
the increased focus, among scientists and portions of the public, on conservation of
biodiversity: (i) – “the accumulation of enough data on deforestation, species
extinction and tropical biology”; (ii) – “the growing awareness of the close linkage
between conservation of biodiversity and economic development”. With the feeling
that a richness, which was unimaginable only a few years before, could be widely
destroyed within a few decades, a radical change occurred. This was remarkably
expressed by Terry L. Erwin (1988, p. 127) : “[…] we should think in terms of more
than 30 million, or perhaps 50 million or more, species of insects on Earth.[…]. The
extermination of 50% or more of the fauna and flora would mean that our genera-
tion will participate in an extinction process involving perhaps 20–30 million species.
We are not talking about a few endangered species listed in the Red Data books
[…]. No matter what the number we are talking about, whether 1 million or
20 million, it is massive destruction of the biological richness of Earth”.
Edward O. Wilson (1988b) gave no precise scientific definition of “Biodiversity”,
but simply stated (p. 3) that: “Biological diversity must be treated more seriously as a
global resource, to be indexed, used, and above all, preserved”. This is symptomatic of
a conservationist and taxonomist conception, regarding biodiversity as a global collec-
tion of species and genes. The defenders of this approach missed the opportunity of
giving genuine scientific meaning to the concept of biodiversity. After 1988, efforts
were made to elaborate scientific definitions, even by conservationist circles; for
example, Jeffrey A. McNeely et al. (1990, p. 17) made the following proposal:
“Biological diversity” encompasses all species of plants, animals and micro-organisms
and the ecosystems and ecological processes of which they are parts. It is an umbrella
term for the degree of nature’s variety, including both the number and frequency of
ecosystems, species or genes in a given assemblage. It is usually considered at three
different levels: genetic diversity, species diversity, and ecosystem diversity”.
In 1994, the International Union for Biological Sciences (IUBS) organized in Paris
an International Forum called “Biodiversity, Science and Development”. In their intro-
duction, Francesco di Castri and Talal Younès (1996) tried to provide a rigorous
definition of biodiversity. Considering the “three levels approach”, they pointed out the
fact that previous definitions of biodiversity paid “little attention, if any, to the
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P. Blandin
interactions within, between and among the various levels of biodiversity” (p. 1). Stressing
the fact that “interaction is the main intrinsic mechanism to shape the characteristics and
the functioning of biodiversity” (p. 2), and considering that the interactions between the
three levels of biodiversity are of a hierarchical nature, forming the “unique trilogy of
biodiversity” (p. 3), they called for a general theory and for the development of a trans-
disciplinary scientific field. Consequently, they proposed a hierarchical definition:
“A more sophisticated definition of biodiversity could be, therefore, the ensemble and
the hierarchical interactions of the genetic, taxonomic and ecological scales of organiza-
tion, at different levels of integration” (p. 4). Moreover, they considered that: “from a
practical viewpoint, structural and functional attributes of system stability, productivity
and sustainability, as well as patterns of ecosystem functioning […], can only be clari-
fied if hierarchies and scales are considered in terms of their interactions” (p. 5). They
concluded that: “The real challenge lies in the possibility of taking into account the
emerging properties that appear by the interactions of the three diversities” (p. 9).
In this way, di Castri and Younès adopted a resolutely holistic approach. They
considered evident the existence of emerging properties, resulting from the hierarchic
organization of biological systems. For example, they considered that the major
attributes of ecosystems (stability, productivity and sustainability) can be under-
stood only by taking into account interactions between the different levels of
integration. Thus, from an epistemological point of view, this functionalist approach
proposes a scientific challenge which is much more stimulating than the taxonomists’
claim to the total inventory of all forms of life.
Nevertheless, the risk still exists that the functionalist approach supports a static
view of a balanced Nature. Demonstrating that we actually have to understand a
“Non-Equilibrium World”, John A. Wiens (1984, p. 440) made the following state-
ment: “Ecology has a long history of presuming that natural systems are orderly
and equilibrial (the “balance of nature” notion; […]), and the infusion of evolutionary
thinking into ecology strengthened this view, providing a mechanism (natural selec-
tion) that may lead to the development of optimally structured communities”. It
may appear paradoxical that an evolutionary thinking favoured the idea of an
Equilibrium Word, but it is a fact that the underlying Balance of Nature Ideology
supported the view of the “climax” ecosystem as a regulated, stable system, opti-
mally composed by a characteristic assemblage of co-adapted species, and showing
the same organization – therefore being homogeneous – everywhere within its
boundaries. Therefore, spatial and temporal heterogeneity has been for a long time
rather an obstacle than a topic for ecological research.
The Decline of the Equilibrium World
The idea of a global “Balance of Nature”, of an “Equilibrium World”, has probably
not only philosophical, but also psychological roots. They could explain the success of
the stability-diversity hypothesis which, as suggested by Goodman (1975, p. 261),
“may have caught the lay conservationists’ fancy, not for the allure of its scientific
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17 Ecology and Biodiversity at the Beginning of the 21st Century
embellishments, but for the more basic appeal of its underlying metaphor. It is the
sort of thing that people like, and want, to believe”. In the context of this ideology,
perturbations were generally considered as catastrophic events.
Landscape ecology favoured an important conceptual shift, introducing a new
way of looking at the spatial organization and dynamics of ecological systems.
Heterogeneity was then considered as an attribute of these systems, and disturbance
as the driving process producing mosaic landscapes (Blondel 1995). Moreover, it
became obvious that any region is characterized by a specific disturbance regime.
These ideas took shape progressively, with such papers as those by Loucks (1970),
White (1979) or Sousa (1984), and the book “The Ecology of Natural Disturbance
and Patch Dynamics” (Pickett and White 1985). Relationships between distur-
bance, patch formation, community structure and species diversity were also
addressed, for example, by Levin and Paine (1974) or Sousa (1979).
At the landscape level, recurring perturbations result in a patchy structure, where
ecological units at different successional states coexist. Consequently, the ecological
diversity and the whole species diversity associated with the landscape are sup-
posed to be sustained by the disturbance regime. Therefore, the mosaic landscape
was substituted for the climatic ecosystem as the effective equilibrium system,
called “metaclimax”, the dynamics of which, driven by the regional disturbance
regime, was recognized as the sustaining process for the regional biodiversity
(Blondel 1986). This shift from “ecosystem equilibrium” to “landscape equilib-
rium” was important, as it allowed spatial and temporal heterogeneity to gain a
conceptual status. But the Balance of Nature Ideology remained in the background,
as it is suggested by the creation of the “metaclimax” concept. That’s why the state-
ment made by Baker (1995, p. 157), on the basis of simulation studies, is of major
interest: “Landscapes with long rotation times may be in perpetual disequilibrium
with their disturbance regimes, because climatic change may create a new distur-
bance regime before the landscape has fully adjusted to the old regime”. Thus,
different landscapes, following different trajectories, should be in different situa-
tions, and may be arrayed along a gradient of states ranging from non-equilibrium
to equilibrium.
In such a new context, the interpretation of species diversity of ecosystems has
to change. Around the 1960s–1980s, under the umbrella of the “equilibrium com-
petitive community paradigm” (Wiens 1984, p. 456), many ecologists considered
that the diversity of a local community is more or less fixed at a level – the satura-
tion point – above which the addition of immigrant species is balanced by the
extinction of pre-existing ones. As stated by Robert E. Ricklefs (1987): “Present-
day ecological investigations are largely founded on the premise that local diversity –
the number of species living in a small, ecologically homogeneous area – is the
deterministic outcome of local processes within the biological community” (p. 167).
But Ricklefs also observed that: “Ecologists are beginning to realize that local
diversity bears the imprint of such global processes as dispersal and species production
and of unique historical circumstance. These processes pose a challenge to community
ecologists to expand the geographical and historical scope of their concepts and
investigations” (p. 167). Ricklefs emphasized the necessity to consider the balance
210
P. Blandin
between local and regional processes, as well as the balance between short term
events and long term processes, to understand the species diversity on the local
scale: “The presence or absence of a species depends on the outcome of processes
tending to increase or decrease its numbers. The latter are generally local in nature
[…]. Most interactions between species are antagonistic, and selection favors
increased competitive ability and predator efficiency. Thus, evolution, while fostering
greater accommodation among coexisting species, ultimately tends to reduce
species richness. Balancing these negative factors is […] the immigration of indi-
viduals from other areas. The variety of immigrants to a particular place depends
on such regional processes as the generation and dispersal of new species (specia-
tion) and also on historical accidents and circumstances related to past climate
history and geographical position of dispersal barriers and corridors. The stronger
speciation and dispersal are, relative to local factors influencing adjustment of
population size and adaptation of individuals, the deeper the imprint of history and
geography on the local community” (p. 169). Moreover, Ricklefs underlined the
fact that the historical dimension of any ecological system results in the diversity
of local situations.
Therefore, everywhere, biodiversity is a stage of “a unique evolutionary play”
(Ghilarov 2000, p. 411). In this perspective, the local species diversity can’t be
considered only as the result of past evolutionary processes, but also as a potential
for further evolution. As early as 1959, Hutchinson suggested that a diversified
community would have a higher aptitude to evolve than a community including a
low number of species: doing so, he introduced the idea that not only the stability
of an ecosystem, but also its adaptability, could depend on its species diversity.
Blandin et al. (1976) developed such ideas. They proposed to consider, schemati-
cally, two different adaptive strategies for ecosystems. On one hand, the adaptability
of ecosystems with low specific diversity would depend on the genetic diversity –
and consequently on the adaptability – of a few species carrying out keystone functions.
On the other hand, the adaptability of ecosystems with high specific diversity
would depend on the existence of functionally redundant species with different
ecological aptitudes. The more numerous are the coexisting species, the lower the
individual number per species may be: within an ecosystem, the genetic diversity –
and, therefore, the adaptability – of each species population can not be independent
of the number of species sharing space and/or trophic resources. Obviously, the two
ecosystem strategies are not exclusive, and any intermediate situation may exist,
depending on the actual number of coexisting species, with a particular hierarchical
interaction between genetic diversity and specific diversity (Blandin 1980).
Similar ideas have been expressed more recently. For example, Tisdell (1995)
compared tropical ecosystems, owning rich biodiversity but being at considerable risk
with any environmental change (because many species have little biological tolerance
and little mobility), with temperate ecosystems showing less biodiversity but species
with greater tolerance and mobility. He concluded that biodiversity as such is neither
necessary nor sufficient to ensure the sustainability of ecosystems. In a more precise
way, di Castri and Younès (1996, p. 5), underlined the possible role of redundant spe-
cies, and the balance between species diversity and genetic diversity: “Not all
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17 Ecology and Biodiversity at the Beginning of the 21st Century
species are equal when it comes to measuring the biodiversity of a system: a few
species can play a keystone role in system functioning, while others may be redun-
dant; some species are dominant and can embrace a very large number of individuals,
thus decreasing the equitability of the system, and others, rare species, may be present
in a very low number of individuals. Also, a lower number of species can be compen-
sated – to a certain extent – by a very high genetic variability in some populations”.
Undoubtedly, the issue of species diversity and genetic diversity interdepen-
dence should help to overcome the controversial hypothesis of a direct relationship
between species diversity and the stability of ecosystems. Moreover, it allows us to
address the question of a community’s capacity to evolve, even if the “Life
Paradox” – life continuity resulting from life change – remains difficult to over-
come. Actually, in the perspective of conservation debates, it is not useful, it is
necessary to consider that evolution is the background and the horizon.
Biodiversity Dynamics: Towards a New Paradigm
Francesco di Castri and Talal Younès (1996, p. 3) called for “a general theory inte-
grating the hierarchical levels of biodiversity, how they come to be and interact”.
The consideration of the relative roles of global, regional and local processes in a
historical perspective, the emphasis on heterogeneity on different spatial and tem-
poral scales, the evidence of various patterns of interdependence between species
diversity and genetic diversity – resulting from history and present ecological con-
texts – provide a possible framework for such a theory, which can be outlined as
follows (Blandin 2004).
The ecosphere’s trajectory is chaotic. At any time the future is unpredictable, at
least for the long term, but the past can be explained by deterministic processes. An
interacting web of global and local processes continuously produces new species
and provokes the extinction of others. At the Earth scale, the balance between species
origination and species extinction processes results in the global dynamics of
biodiversity.
The number of species living within a region results, on one hand, from extra-
regional species originations and subsequent immigrations, and from intra-regional
species originations. On the other hand, these processes are counterbalanced by
intra-regional extinctions and the shifting of species out of the region, for example
in relation with climatic changes.
Within a regional heterogeneous landscape, the richness of a local assemblage
of species depends first on the regional stock of the species which are capable of
coexisting. Secondly, it depends on the landscape structure, which governs the
migration flows between ecological units. The actual coexistence of populations of
different species depends on the physical and chemical constraints of the environ-
ment, and on the local flows of resources that can be exploited by these species,
according to their needs. It also depends on the variety and regime of disturbances.
Lastly, catastrophic events may produce dramatic changes.
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P. Blandin
Taking into account these new insights, natural communities can be viewed as
being arrayed along a gradient of situations resulting on one side from a history of
stochastic events, on the other from long environmental stability, with intermediate
situations resulting from more or less long periods characterized by more or less
regular disturbance regimes. According to such different histories, the relative roles
in patterns of selection, of environmental characteristics, of trophic flows and of
inter-specific interactions may have changed widely: biotic coupling, allowing
co-selection, is unlikely under stochastically changing conditions, but could be
possible in recurrent contexts, giving rise to co-adapted species.
In each situation, each species is distributed through the landscape mosaic in a
particular way, and participates in assemblages of species interacting in a more or
less strict manner, according to the degree of co-adaptation reached through evolution.
The genetic diversity of each species has a particular pattern, depending on the
number and size of populations, on the local flows of individuals between popula-
tions and on the balance between immigration and emigration. The local sustain-
ability of the species depends on this genetic diversity. The size of each species
population is partially determined by the number and size of the populations of
other species sharing space and resources: there are necessary interactions between
the genetic diversity of species and the species diversity of assemblages, in the
framework of the local ecological diversity. Therefore, the adaptative capacities of
the biotic communities – on which depends their sustainability – are governed by
the present pattern of interaction between the three biodiversity levels.
At any time, on any spatial scale, the global biodiversity is the unique heritage of
past evolution, and the unique, limited potential for further evolution. At any time,
many different trajectories are possible, but only one will be followed. As local,
regional and global processes are continuously interacting, the evolution of ecological
systems must be considered as a “web of interdependent trajectories”: what we can
call the “Transactional Trajectories Paradigm” – using “transactional” in the spirit of
Dewey’s epistemology (Dewey and Bentley 1949) – is taking the place of the
Odumian Ecosystem Paradigm, deep-rooted in the Balance of Nature Ideology.
The Sustainable Adaptability of the Ecosphere
Evolution cannot be considered only in the limited sense of species originations and
extinctions: it is a global process of ecological changes, co-evolutionary interactions
and biodiversity transformation. Broadening the perspective at the Ecosphere level,
the Israeli ecologist Zev Naveh (2000) has suggested that “The Total Human Eco-
system should be regarded as the highest coevolutionary ecological entity on Earth”
and he maintained that “This conceptual approach enables us to view the evolution
of Total Human Ecosystem landscapes in the light of new holistic and transdisci-
plinary insights into the dynamic process of self-organization and coevolution in
nature and human societies” (p. 358). Then, the Life Paradox could be overcome
saying that coevolution makes possible life’s sustainability, with new species substituting
213
References
Baker WL (1995) Longterm response of disturbance landscapes to human intervention and global
change. Landscape Ecol 10(3):143–159
Bergandi D (1995) ‘Reductionist holism’: an oxymoron or a philosophical chimaera of E.P.
Odum’s systems ecology. Ludus Vitalis 3(5):145–180, reprinted in Keller, D.R. & F.B. Golley
(eds.) 2000. The philosophy of ecology: from science to synthesis (abridged version),
University of Georgia Press, Athens, pp 204–217
Blandin P (1980) Evolution des écosystèmes et stratégies cénotiques. In: Barbault R, Blandin P,
Meyer JA (eds) Recherches d’écologie théorique. Les stratégies adaptatives. Maloine, Paris,
pp 221–235
Blandin P (2004) Biodiversity, between science and ethics. In: Hanna S, Mikhail WZA (eds) Soil
zoology for sustainable development in the 21st century. Egypte, Cairo, pp 3–35
Blandin P, Lecordier C, Barbault R (1976) Réflexions sur la notion d’écosystème: Le concept de
stratégie cénotique. Ecol Bull 7:391–410
Blondel J (1986) Biogéographie évolutive. Masson, Paris
Blondel J (1995) Biogéographie. Approche écologique et évolutive. Masson, Paris
Dewey J, Bentley AF (1949) Knowing and the known. Beacon, Boston
others to perform continuously ecological functioning. This implies an ideological
shift, “sustainability” being substituted for “conservation” as the fundamental
aim.
This evolutionary perspective opens new insights. As the unique memory of the
past evolution, with the scattered remains of the paleobiodiversity, the present
biodiversity offers keys for understanding evolutionary processes, as well as oppor-
tunities for wonder. With the Russian ecologist Alexei M. Ghilarov (2000), we can
say that it is “an example of evolutionary heritage that is probably worth protection
no less than the heritage of our culture” (p. 411). Moreover, as the unique potential
for further evolution, the present biodiversity is the unique available man’s companion
for the co-evolutionary adventure to come. To satisfy, through the generations,
humans’ needs implies the permanent availability of natural “resources” able to
match the cultural diversity of humans’ desires. Today, humans’ aims and projects
are diverse; tomorrow, they will be different; nobody can predict what future
generations, living in new ecological and cultural contexts, will need and desire.
Therefore, the main problem is to transmit, through generations, a “four-levels
diversity” allowing the sustainability of “man-nature systems”: this implies not
only functional continuity, but also the maintenance of a capacity to evolve, which
depends on the integration of climatic, specific, genetic and cultural diversities
within man-nature systems. Obviously, research is needed to provide new insights
into the interactions of these four diversities, on all spatial scales. Thus, no longer
can Ecology be merely a “biological” or a “natural” science: Ecology must evolve
in order to contribute to the construction of a transdisciplinary field, dealing with
integrated natural and socio-cultural trajectories. This is necessary to provide
the scientific bases that humans need to ensure the “sustainable adaptability of the
Total Human Ecosystem”.
17 Ecology and Biodiversity at the Beginning of the 21st Century
214
di Castri F, Younès T (1996) Introduction: Biodiversity, the Emergence of a New Scientific
Field–Its Perspectives and Constraints. In: di Castri F, Younès T (eds) Biodiversity, science
and development. Towards a new partnership. CAB International and IUBS, Paris, pp 1–11
Erwin TL (1988) The tropical forest canopy. The heart of biotic diversity. In: Wilson EO, Peter
FM (eds) Biodiversity. National Academy Press, Washington, DC, pp 123–129
Ghilarov AM (2000) Ecosystem functioning and intrinsic value of biodiversity. Oikos 90:408–412
Golley FB (1991) The ecosystem concept: a search for order. Ecol Res 6:129–138
Goodman D (1975) The theory of diversity-stability relationships in ecology. Q Rev Biol
50(3):237–266
Harroy JP (1949) Définition de la protection de la nature. In: UIPN (ed) Documents préparatoires
à la conférence technique internationale pour la protection de la nature. UNESCO, Paris,
Bruxelles, pp 9–14
Hutchinson GE (1959) Homage to Santa Rosalia, or why are there so many kinds of animals? Am
Nat 93:145–159
Leigh EG (1965) On the relationship between productivity, biomass, diversity, and stability of a
community. Proc Natl Acad Sci USA 53:777–783
Levin SA, Paine RT (1974) Disturbance, patch formation, and community structure. Proc Natl
Acad Sci USA 71:2744–2747
Loucks OL (1970) Evolution of diversity, efficiency and community stability. Am Zool 10:17–25
Macfadyen A (1975) Some thoughts on the behaviour of ecologists. J Anim Ecol 44:351–363
Margalef R (1969) Diversity and stability: a practical proposal and a model of interdependence.
Brookhaven Symp Biol 22:25–37
McNeely JF et al (1990) Conserving the world’s biological diversity. IUCN/WRI, CI, WWF-US,
The World Bank, Gland, Switzerland, Washington, DC
Naveh Z (2000) The total human ecosystem: integrating ecology and economics. Bioscience
50(4):357–361
Odum EP (1953) Fundamentals of ecology. W.B. Saunders, Philadelphia
Paine RT (1966) Food web complexity and species diversity. Am Nat 100:65–75
Petit G (1950) Protection de la nature et écologie. In: IUPN (ed) International technical conference
on the protection of nature, Lake Success, 22-29-VIII-1949, proceedings and papers.
UNESCO, Paris, Bruxelles, pp 304–314
Pickett STA, White PS (eds) (1985) The ecology of natural disturbance and patch dynamics.
Academic, New York
Ricklefs RE (1987) Community diversity: relative roles of local and regional processes. Science
235:167–171
Smith FE (1972) Spatial heterogeneity, stability and diversity in ecosystems. Trans Conn Acad
Arts Sci 44:309–335
Sousa WP (1979) Disturbance in marine intertidal boulder fields: the nonequilibrium maintenance
of species diversity. Ecology 60:1225–1239
Sousa WP (1984) The role of disturbance in natural communities. Annu Rev Ecol Syst
15:353–391
Tansley AG (1935) The use and abuse of vegetational concepts and terms. Ecology 16:284–307
Tisdell CA (1995) Issues in biodiversity conservation including the role of local communities.
Environ Conserv 22(3):216–222
UIPN (1948) Union internationale pour la protection de la nature, créée à Fontainebleau le 5
octobre 1948. UIPN, Bruxelles
White PS (1979) Pattern, process and natural disturbance in vegetation. Bot Rev 45:229–299
Wiens JA (1984) On understanting a non-equilibrium world: myth and reality in community patterns and
processes. In: Strong DR Jr, Simberloff D, Abele LG, Thistle AB (eds) Ecological communities:
conceptual issues and the evidence. Princeton University Press Princeton, New York, pp 439–457
Wilson EO (1988a) Biodiversity. National Academy Press, Washington, DC, pp v–vii
Wilson EO (1988b) Biodiversity. National Academy Press, Washington, DC, pp 3–18
P. Blandin
215
Chapter 18
An Ecosystem View into the Twenty-first Century
Wolfgang Haber
Introduction: Ecosystem Between Recognition and Disputation
The term “ecosystem”, introduced by Arthur G. Tansley1 in his epoch-making article
of 1935, has provided us with a useful and promising concept to be applied for inves-
tigative understanding and solving the growing environmental problems of the twenty-
first century. For both institutions and persons bearing responsibility for a sustainable
development, usage of “ecosystem management” and “ecosystem services” has
become their common language and means of communication. The “Millennium
Ecosystem Assessment” launched by the United Nations at the turn of the millennia
(MEA 2003) will further strengthen general attention for that rather abstract term and
turn it into a self-evident component of the twenty-first century’s everyday publicity.
Among environmental scientists, however, notably in the disciplines of ecology,
biology, and geography, and their theorists and practitioners, “ecosystem” has been,
and is being debated as a more or less contentious term. In order to bridge upcoming
differences between scientific rigour and clarity on one side, and general, transdis-
ciplinary usage of “ecosystem” in public affairs on the other, and to avoid further
misunderstandings and errors, it appears useful to take a retrospective view of the
history of the term and its varying meanings in theory and practice (cf. Allen et al.
2005; Peterson 2005; de Laplante 2005; Blandin 2006).
“Ecosystem” was the clear winner when in 1988, at the occasion of the 75 years’
jubilee of the British Ecological Society the members were asked about the most
important concepts in ecology. It was followed, but at a considerable distance, by
“succession” (Cherrett 1989). “Ecosystem ecology” had become established as a
1 Willis 1997 has pointed out that the term was originally coined by Clapham whom Tansley had
asked “if he could think of a suitable word to denote the physical and biological components of
an environment considered in relation to each other as a unit. When Clapham suggested ‘ecosystem’,
Tansley, after some consideration, wholly approved of it” (p. 268). In his article of 1935, however,
Tansley did not mention Clapham.
W. Haber (*)
Technische Universität München, WZW, Lehrstuhl fur Landschaftsökologie,
Emil-Ramann-Strasse 6, D-85354 Freising, Germany
e-mail: WETHABER@aol.com
A. Schwarz and K. Jax (eds.), Ecology Revisited: Reflecting on Concepts, Advancing Science,
DOI 10.1007/978-90-481-9744-6_18, © Springer Science+Business Media B.V. 2011
216
W. Haber
special branch of ecology (Pomeroy and Alberts 1988), and the popularity both of
the term and the concept “ecosystem” literally reached their peak. E. O. Wilson
(1996) even counted the ecosystem among the “metaphysical constructs” that would
have proven more powerful and less vulnerable than ordinary scientific theories.
Since 1998 it can also boast a special scientific periodical entitled “Ecosystems”.
This seemed to be a triumphant success for the term Tansley had published – but
not precisely defined–about 50 years earlier. Stations of this success were
Lindeman’s (1942) pioneering study of a small lake, and in particular E. P. Odum’s
ground-breaking textbook “Fundamentals of ecology” (1953) which was based on
the ecosystem as its central concept; Bergandi (1995) called it one of the few really
paradigmatic scientific texts. The ecosystem then became the core concept of the
first great international research programmes launched since the 1960s, such as the
International Biological Programme (IBP) and its follower “Man and the Biosphere”
(MAB). Their worldwide success and reputation considerably strengthened and
popularized the use of the ecosystem concept and were unthinkable without it.
In contrast to this success story, since the 1980s a growing number of ecologists
began questioning the primacy of the ecosystem concept and started a controversial
debate on its meaning and importance (cf. Reiners 1986, Likens 1992). In particular
evolutionary and population biologists developed an aversion against ecosystem
studies to which they reproached one-sided consideration of energy and matter
transfers or trophic levels, neglecting or ignoring aspects of population and community
ecology as well as biodiversity. In Holling’s (1992, 1996) view, population and
community ecology seemed to exist in their own world and ecosystem ecology in
another one. O’Neill et al. (1986) confronted them, emphasizing that an integral
view of an ecosystem requires a combination of the two approaches, because the
many functions of an ecosystem cannot be understood without considering biotic
interactions in all their variety.
As a matter of fact, E. P. Odum, the father of ecosystem ecology, did take well
into account population and community ecology, which occupy special and impor-
tant chapters both in the three editions of his above-mentioned textbook and in the
various popular versions following it. Nevertheless, Bergandi (1995) called the
Odumian ecosystem approach utterly reductionist. This holds, however, rather for
the work of Howard T. Odum, Eugene’s brother, who factually reduced all ecosystem
processes to energy transformations. Eugene Odum, it is true, recognized and supported
his brother’s view, incorporating it into his own books, but always insisted upon his
(Eugene’s) approach as being truly “holistic”.
Ecosystem Management and Ecosystem Services
The power of the ecosystem concept proved itself anew in America when in 1992
the U.S. administration, following a proposal of the then Vice President Gore,
decided to change the management of the country’s natural resources from a single-
resource approach (forests, rangeland, wildlife etc.) to a comprehensive “wholistic”
one for which the name “ecosystem management” was chosen. Although ecosystems
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as subjects of science were controversially debated among ecological scientists, they
appeared to be rather easily utilized as subjects of environmental management.
About 20 U.S. federal agencies formed an “Interagency Ecosystem Management
Task Force” which in 1995 presented a comprehensive three-volume report entitled
“Healthy ecosystems and sustainable economies − the Federal Interagency
Ecosystem Management Initiative” (EMI). Its goal was a proactive approach to
ensuring a sustainable economy and a sustainable environment through ecosystem
management (Malone 1995). This initiative may be considered as important as that
of the Biodiversity Convention of 1992 embracing ecosystem diversity–albeit on
a national scale.
The EMI, however, enhanced the confusion about the ecosystem concept
(Carpenter 1995). To cite from Stein and Gelburd (1998, p. 74; emphasis WH):
The ecosystem approach is a method for sustaining or restoring natural systems and their
functions and values. It is based on a collaboratively developed vision of desired future
conditions that integrates ecological, economic, and social factors and is applied within a
geographic framework defined primarily by ecological boundaries. The ecosystem
approach was chosen to restore and sustain [reversed priority! W.H.] the health, productivity,
and biological diversity of ecosystems and the overall quality of life – through a natural
resource management approach that is fully integrated with social and economic goals.
This somewhat curious statement leaves one wondering whether the ecosystem
approach is restricted to natural resources or natural systems, which is justified
because ecosystem management per se does not include economic and social factors,
or should include these aspects. Referring to the EMI, Szaro et al. broaden its
vagueness by stating: “For practical purposes, ecosystem management is generally
synonymous with sustainable development, sustainable management … and a number
of other terms being used to identify an ecological approach to land and resource
management” (1998, p. 5).
The same authors state a few paragraphs further on that “[the] ecosystem approach
emphasizes place- or region-based objectives” because ecosystem data are “inher-
ently spatial” (Brussard et al. 1998), and that “Ecosystem planning must consider the
dynamics of landscape scale patterns” (Szaro et al. 1998, pp. 2 f.). The seven case
studies forming part of the EMI report included among others a large watershed, the
Great Lakes basin, the whole of South Florida, and the Prince William Sound in
Alaska (which had been devastated by the wreckage of the oil tanker “Exxon
Valdez”). It seems that “ecosystem” was used here as a kind of umbrella term for
quite a variety of management goals and measures, comparable to the German
(untranslatable) term “Naturhaushalt”. In a commentary on EMI, Sheifer (1996) con-
sistently used the double term “ecosystems/ecoregions”, suggesting a regional land-
scape dimension of ecosystem (cf. Blandin and Lamotte 1988).
The EMI, however, was not allowed to fulfil its objectives – not because of the
confusion about the ecosystem concept, but for political reasons: the Republican
majority in the US Congress, outcome of the 1996 federal elections, rejected the
initiative and cut the funds for it. Nevertheless, the journal “Landscape and Urban
Planning” devoted a 234-page special issue (Vol. 40, 1998) to the EMI which offers
interesting reading for ecosystem discussions. Enhancing the confusion, a book
entitled “Ecosystem Management”, edited by the (American) authors Samson and
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W. Haber
Knopf (1996), did not mention EMI in its final report at all but wrote in the preface
“The fact that no one knows what ecosystem management means has not diminished
enthusiasm for the concept” (Wilcove and Samson 1987, p. 322). And in 1995 the
Ecological Society of America issued a document entitled “The scientific basis for
ecosystem management”, again without mentioning EMI – and as such criticized
by Zeide (1996) for not being as scientific as intimated. A number of ecologists and
other experts who had cooperated in the EMI worked out the results and put them
together in a three-volume book entitled “Ecological stewardship: A common refer-
ence for ecosystem management” (Johnson et al. 1999). The emphasis had shifted
from management to stewardship, with a bias towards both conservation biology
and the biological features of the ecosystem, thus reflecting the growing interest in
biodiversity.
Towards the end of the twentieth century, the notion of “ecosystem services”
offered to people and to society in general attracted great public attention, in
particular stimulated by Daily’s (1997) book. These services linked ecosystems to
what ecological economists had conceived of as “natural capital” in the then
upcoming debate on “sustainable development”. It is these ecosystem services
which have served as the starting point for the newest and comprehensive global
environmental programme called “Millennium Ecosystem Assessment” which, as
mentioned at the beginning, was launched by the United Nations in June 2001
(MEA 2003). This scientific enterprise focuses on how changes in ecosystem
services have affected and are affecting human well-beings, how ecosystem
changes may affect people in future decades, and what types of response can be
adopted at local, national, or global scales to improve ecosystem management and
thereby contribute to human well-being and poverty alleviation. Ecosystem services,
however, are not based on clearly delimitable environmental objects, but on func-
tional units of variable size and composition. Regarding ecosystem management or
stewardship, we are faced with a big problem of implementation requiring immense
mental effort which is based on the ecosystem concept – both loading it with enormous
expectations and implications and imposing a likewise huge responsibility on
ecological science to develop a theoretically and practically reliable basis for the
concept. This is clearly demonstrated in the (preliminary) synthesis report titled
“Ecosystems and Human Well-Being” (MEA 2005).
The Ecosystem’s Position in Hierarchies and Scales
How well are ecologists prepared to scientifically accompany, advise on and
support the solving of this huge task of the twenty-first century? After all the
controversies of the last decades of the twentieth century Mayr (1997) had
argued that the ecosystem concept – after its great Odum-inspired popularity in
the 1960s and 1970s – had lost its role of a dominant paradigm. O’Neill (2001),
one of the protagonists of ecosystem ecology, had even asked “Is it time to bury
the ecosystem concept? (With full military honours, of course!)”, but gave the
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answer himself: “Probably not”. The term which thinkers had earlier referred to as
an epistemological nightmare, quite contrary to the view of Scheiner et al. (1993),
has obviously become indispensable. How should it be handled? Again, a short
retrospective appears helpful for finding an answer.
When Clapham suggested the term “ecosystem” to Tansley in the early 1930s
(Willis 1997), he borrowed the word “system” from the then upcoming system
studies. Of course “system” implies a holistic view, but at the same time empha-
sizes a physical character and a mechanical or machine-like approach – an analogy
that, by the way, had appeared much earlier (1922) in Tansley’s work. According
to Golley (1993) and Jax (2002), the main motivation of Tansley, whom Evans
(1976) and quite recently Sheail (2005) characterize as continuously preoccupied
with philosophy and processes of thought, was to strengthen the recognition of
ecology as a serious science and to keep it free from misleading philosophical
abuses of holistic theories of his time.
There are two important aspects in Tansley’s article that should be emphasized.
The first is that he explicitly called the ecosystem a “mental isolate”, i.e. a “meta-
physical construct” to put it into E.O. Wilson’s terms. The second one is that Tansley
conceived of a sequence of organization levels of nature – he called them “physical
systems of the universe” – “which range from the universe as a whole [!] down to the
atom” (Tansley 1935, p. 299), and that he assigned to the ecosystem a specific level
within this hierarchical sequence. It is astonishing that this second, no less important
concept (Schultz (1967) even called it “far more important”) has met with relatively
little appreciation until today. It was Egler (1942) who took up this idea for his vegetation
system and formulated a first but still incomplete hierarchical sequence of levels of
organization of living nature, which was complemented by Novikoff (1945).
Feibleman (1954) even propounded a “theory of integrative levels” based on twelve
“laws”, which were discussed in an ecosystem context by Schultz (1967). (These
three authors, by the way, are never cited in more recent publications on hierarchy.)
Tansley’s idea of the ecosystem as a mental construct means that the object does
not exist as such in reality, but only in the researcher’s mind, and it is here where it
is defined regarding its content and especially its boundaries. This mental picture is
then projected onto a real situation in nature, the mental boundaries being fitted to
factual ones such as river banks and lake shores, forest edges, watershed boundaries
or boundaries originating from substrate or soil differences, all of which can serve
to delimit ecosystems. In today’s landscapes, however, most boundaries result from
human land use, although they may coincide with natural boundaries. Clearly, if the
ecosystem is but a mental construct, its boundaries are set by the observer or inves-
tigator! Therefore, to liken an ecosystem to a Clementsian (super)organism is
fundamentally wrong and misleading, because an organism has an outside boundary
(skin, epidermis, cell wall etc.) produced by itself and accepted by the researcher.
Tansley’s original concept – it was really only an idea – of the hierarchy of physical
systems in nature ranked these from the smallest dimension to the largest, and for
him the ecosystem was but one of these systems, apparently of an intermediate
dimension (a “middle-number system” sensu Weinberg 1975). But this dimension
must not be confounded with spatial scale, which would imply that, according to its
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W. Haber
position in the hierarchy, an ecosystem is on a larger spatial scale than a community
or a population and should be treated accordingly – which lacks any logic. Moreover,
many biotic processes like succession or migration occur on larger spatial or temporal
scales than ecosystem processes such as nutrient cycling.
Roots of Confusion
In this context, we hit upon one of the roots of misunderstandings, semantic confusions,
and contradictions encumbering the ecosystem concept. Lindeman’s pioneering
ecosystem study of a small lake (only 1 ha surface, 1 m deep) had assigned an
explicit spatial dimension and a clear boundary (the shoreline) to Tansley’s concept,
so it fitted into his hierarchical order. Eugene Odum, however, in his 1953 book
presented the ecosystem as a basic unit in ecology of every spatial dimension, from
the population to the biosphere, a definition which was taken up and confirmed by
Evans (1956). But in the second edition of his book (1959), Odum followed
Tansley, restricting the ecosystem to a specific level in the hierarchy – but continued
nevertheless to use the term in its broader sense too, and nobody appeared unduly
disturbed by these contradictions (Golley 1993, p. 72).
This explains the ongoing confusion concerning the character of the levels in
Tansley’s (and his followers’) hierarchy, i.e. between a scalar (spatial) level and an
organizational level – which induced Wiegleb (1996) and Jax (2002), following
Allen and Hoekstra (1990), to suggest replacing “organizational” with “observational”.
It also resulted in a proliferation of “branched” or “broken” hierarchies (see the
discussion in Wiegleb 1996, chapter 3.4) to avoid this confusion. The book by
Hölter (2002) may contribute to its elucidation. Anyway, the validity of hierarchy
theory above the organism level – where all levels are “mental constructs” – is more
strongly contested than at distinctly lower levels of organization (Fränzle 2001, p. 75).
Thus the concept of ecosystem in the traditional hierarchy scheme runs counter to
the concept of ecosystem as a perspective that can be studied as a whole, incorpo-
rating activities from different scales (Vogt et al. 1996).
A solution of this scale-related dilemma was proposed by Ellenberg (1973; see
also Mueller-Dombois and Ellenberg 2002, p. 168–171) who, following Evans’
ecosystem definition in principle, established a corresponding “system of ecosystems”.
He distinguished five classes of ecosystems ordered in a hierarchy of spatial dimen-
sions, namely mega-, macro-, meso-, micro- and nano-ecosystems. He called the
meso-ecosystems the “ecosystems in the strictest sense” (Ökoysteme im engeren
Sinn, p. 237) and the basic types of ecosystem classification, thus approaching once
again the ecosystem position in Tansley’s (and the later Odum’s) hierarchy. An
example of a meso-ecosystem is the summer-green broadleaved forest which is
subdivided into the different alliances and associations familiar to phytosociologists.
However, Ellenberg’s classification did not arouse much attention.
Ellenberg’s understanding of ecosystem is reflected in the German IBP project
in the Solling which he directed (Ellenberg et al. 1986) and which is regarded as a
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“milestone in the development of ecosystem research” (Fränzle 1998, p. 11). It was
strongly influenced by the phytosociological approach of the Zurich-Montpellier
school based on plant associations recognized by their floristic composition and
character species. A plant community with its more or less distinct boundaries
could be mapped and rather easily be likened to a (micro-)ecosystem. This equal-
ization (more in practice than in theory) of plant associations or formations with
ecosystems (meso- and micro-ecosystems in Ellenberg’s terminology) is quite
common in continental European ecosystem studies, especially when applied to
nature conservation or landscape planning, and is considered a useful and proven
approach to ecosystem studies. Both nature conservationists and environmentalists
soon adopted the term “ecosystem”, making liberal use of it and opened its way into
everyday language.
It was during the IBP research that, almost imperceptibly, the “ecosystem”
changed from Tansley’s “mental isolate” to a real object in nature or, as Schultz
(1967) put it, a “perceptible object of study”. Thus ecosystem ecology owed its
most important success to “a realist, even ontic perspective of the ecosystem as a
real world entity” (Potthast 2002, p. 139).
The Ecosystem Concept and Theoretical Ecology
It was not until the 1990s that theoretical ecologists seriously tackled the epistemo-
logical background of the ecosystem concept. This Handbook is both a proof and a
first essential result of this endeavour. At first they seemed to be caught in the
“holism trap”. Users of the concept are often criticized for favouring “organicist
thinking” or adhering to Clementsian “superorganism belief” (cf. Trepl 1988, de
Laplante 2005), often voiced with a hidden derogatory undertone. Such critique,
however, is one-sided and leads one astray. No organism can exist without interacting
with other organisms, be it in the form of symbiotic or antagonistic relationships.
These constitute, especially when displaying a recurrent behaviour, a real network
whose structure can be disclosed analytically without any organicist idea in mind,
and which can of course be called a system. Such reasoning is recognized by Trepl
(1988), without however curbing the tendency of lumping together all systems- or
synthesis-minded ecologists into one organicist or “holist” box.
Therefore, ecological scientists (and also many biologists) remain seriously split
on the issue of reductionism versus holism, instead of dealing simultaneously with
the part and the whole (Blandin 2006). A reductionist approach is obligatory in any
investigation of highly complex phenomena, as complexity can only be mastered
by breaking it down into its (presumably) principal components. This analytical
procedure, however, has to be followed, and completed by a re-synthesis of the
components in order to comprehend the functioning of the complex whole or the
“why” of complexity. Such a combination of a reductionist with a holistic approach,
often done iteratively, is essential for ecosystem investigations, but is neglected by
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researchers who shun the “holism trap”. According to Schultz (1967), this attitude
goes back to the origins of science and the historical primacy of physics in the
seventeenth and eighteenth century. The objects of the physicist’s manner of scientific
inquiry were inanimate and timeless matter and energy, enabling him to explain the
universe by means of a purely reductionist approach and to rather easily find
generalizations and laws. This became the “only justifiable” scientific method2
which was imposed on late-comers such as the life sciences; these dealt with bio-
logical entities each of which is unique - an “event” or a “chunk of space-time” that
is only typifiable and defies explanation in purely physical terms. Therefore, biology
is essentially an “event science”; there are only rules, but no laws at the biological
level, theories are essentially non-testable, and the life phenomena too indeterminate
for assignment of causes and predictions. Moreover, one of the fundamental
biological questions, that is even constitutive for ecology, addresses the assessment
of environmental factors with regard to their suitability for, and promotion of, the
self-maintenance of organisms. This is ultimately a teleological question (cf. Weil
2005) greatly exceeding “common” scientific laws and utterly reprehensible, if not
abhorrent for physicists’ minds; but it can be justified by heuristic necessity forgoing
causal explanations. What would have been the development of modern science if
biology instead of physics had achieved primacy? It might have been considerably
delayed, because those early scientists of reductionist obsession would have been
totally overcharged with tackling the huge diversity of life and living phenomena.
Schwarz (2003) has suggested a thoughtful and promising mode to bridge these
abysmal differences. She places ecology as a kind of “third power”, endowed with
mediating abilities, between the two mainstreams of biology: (1) physiology as
reductionist-analytical research pursuing explanatory goals, based on physicist
reasoning and seeking general or generalisable laws, and (2) physiognomy founded
upon integrative or synthetic comprehension (not explanation) of principally unique
phenomena of “Gestalt” quality. Schwarz also gives full recognition to the indis-
pensable importance of metaphors (“metaphysical constructs” sensu E. O. Wilson)
and of heuristics. Such a mediating role of ecology, however, is never uniform, but
varies or fluctuates according to changing preferences for central fundamental con-
cepts of ecology, namely “energy”, “niche” and “microcosm”. The latter might be
understood as “ecosystem” in its narrow definition, so it fits well into Schwarz’s
arguments. These are apt to break the unseemly haughtiness of the reductionist
party and its unfounded superiority based on (mostly unconscious) incontestability
of their scientific attitude.
The power, or pervading influence, of the ecosystem concept appears to be based
on the deep-felt yearning of ecologists for “unifying ideas”, which has continually
been articulated since about 1960–it was the topic of the first International Congress
2Fränzle, however, points out that leading representatives of the scientific community around
1800, like Buffon, James Hutton, Georg Forster, Linné, Miraband, Kant were already primarily
interested in defining interrelationships, and less so “in detailed generic analyses or unravelling
specific microscale causalities” (2001, p. 60).
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18 An Ecosystem View into the Twenty-first Century
of Ecology in The Hague 1974. Ford and Ishii (2001), repeating the plea for a
synthesis in ecology, consider “integrative concepts” an essential part of ecology
because they enable scientists to organize and reorganize ecological knowledge as
the science progresses. Who would contest the integrative notion of the ecosystem
concept? Indeed, the two authors recommend, as a viable way to achieve synthesis,
the construction [!] or development of integrative concepts regarding the internal
organization of ecological systems. Fränzle argues that “the present theoretical
background to ecosystem research has not yet reached the level of a comprehensive
unified theory”, but he sees “a commendable number of unifying concepts and
integrative approaches with regard to ecosystem research” (2001, p. 83). In his
opinion, all these concepts are derived from, or associated with, system theories in
general, more specifically with those of self-organizing ecological systems. The
latter integrate components of thermodynamics, dissipative processes, information
and network theories, game theory, catastrophe theory and hierarchy theory. This
appears once more as a rather “physicalist” way of thinking, to which a physical
geographer like Fränzle is inclined. His opinion, however, that the complexity of
ecosystems can only be expressed in mathematical models, has to be questioned
because these ignore or omit the incalculable behaviour of living agents, leaving the
modeller with incomplete or only partial knowledge of the system which he seeks
to understand. As Schaffer (1985) put it, ecologists will never be able to write down
the complete governing equations for any natural system.
Society’s view of science, it is true, still consists of a trilogy of theories, hypotheses,
and laws to explain observed facts and to allow prediction of events. But in ecology,
this legacy of a reductionist, quantifying scientific problem approach has failed in
the face of the huge complexity of the environment, which is much greater than the
earth’s climate system of which, as we know, only probable futures can be assessed.
Environmental impact assessments (EIA) of projects, which are mandatory in many
countries, require such predictions but they simply cannot be provided with the reli-
ability taken for granted – an example of unqualified and irresponsible legislation.
Outlook: A Plea for Realism and Transdisciplinarity
O’Neill’s (2001) cautious answer to his question of burying the ecosystem concept
read: “Probably not”. All attempts to basically question it, not to mention abolish
it, have failed as yet, proving both the power of the concept and its utility. It appears
that we are unable, even as scientists, to handle complexity (the dominant property
of the biosphere or the natural environment) on the exclusive basis of clear scientific
reasoning utilizing Kuhnian principles and logical theory, without resorting to
heuristics. Thus, refraining from reliable predictions of future environmental states,
we have to search for a general understanding of the complex dynamics of the
environment in order to find approaches for suitable, practicable management
strategies. For this purpose, the ecosystem concept has opened the right pathways,
offering opportunities to approximate, contribute to, or even achieve basic societal
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W. Haber
goals such as sustainable development, environmental health, safety and security,
biodiversity handling as well as general welfare. Accordingly, new scientific
ecological disciplines such as conservation biology, ecological economics, ecological
planning and management are relying on, and supported by ecosystem ecology in
order to increase the controllability of an uncertain world (de Laplante 2005).
In these more applied research fields, ecosystem ecology is becoming an influential,
diversifying branch of ecological science, meeting with growing public attention
and support. Furthermore, it transcends traditional disciplinary boundaries, bridging
the gap separating it from social science and humanities (Cantlon 2002;
Haber 2004). In a time of rapid social and environmental change, such transdiscipli-
narity – a visualisation already emphatically pursued by Eugene Odum–has become
indispensable.
Orthodox scientific circles, however, even in ecology, still frown upon such
“soft” science and its value-laden, issue-driven approaches. Although “hard” reduc-
tionist science based on quantitative mathematical models is proving unable to
devise practicable means for solving complex environmental problems, its episte-
mological scruples should not be dismissed altogether. Ford and Ishii (2001),
despite their strong support for “integrative concepts” in ecology, warned both of
the conjectures encumbering them and of the lack of a recognized procedure for
synthesis. Following Taylor and Haila (2001), such concepts are unstable and elusive
and cannot be taken for granted, so we have to reconceptualise them from time to
time. Even when the ecosystem concept has already proven indispensable for envi-
ronmental understanding and management, we should be aware of such scruples
(particularly of its theoretical weaknesses and “birth defects” mentioned above) and
stick to understanding it as a “metaphysical construct” or metaphor sensu Schwarz
(2003). Whenever we apply this construct to a real piece of nature whereby we
delimit ourselves according to the purpose of ecological research, we have to be
careful that the application is precisely defined, even if ecology still lacks termino-
logical rigour (Breckling and Müller 1997). There will always be a temptation to
slide from the metaphysical construct into tangible reality of nature. But it remains
a helpful and fruitful image.
Ecology in general, and ecosystem ecology in particular, must no longer be
confronted with the dichotomy between reductionism and holism – even less by
insinuating that the latter is scientifically reprehensible. This dichotomy, an application
of Hegelian dialectic, might have served a heuristic purpose, but diverts attention
from the principal goal of ecological research: to explain life in its organization, its
universality as well as its diversity of change, and all of them in their dependence
upon, adaptation to, and limited capability of alteration of the non-living, physico-
chemical environment. The “third way” suggested by Schwarz (2003), mentioned
above, can lead in the right direction, making use of narrative and “good metaphors”
because to use metaphor well is to discern similarities.” (Aristotle in Poetics
1459a: 6–7). In this way, we get back to, and revive the far-sighted and invaluable
legacy of Alexander von Humboldt to science and art, “his sense of unity within the
complexity of what we now call ecosystems” (Fränzle 2001, p. 63).
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3 die Lehre von der in der Allgemeinheit der Gesetze fundierten Einmaligkeit von Situationen
(Picht 1979, p. 25).
It is also appropriate to cite Georg Picht’s (1979) definition of ecology as – “the
science of the singularity of situations, founded on the generality of natural laws”.3
Picht’s reasoning stems from an understanding of life (which he sometimes identified
with “oikos”) as a “contemplative conception of nature, being interpreted in a
normative way” and insists upon this contemplative relationship, with nature being
superior both to the scientific (ecological) and the economic-technical relationship
with nature. The general tendency of science – in the eyes of the public supporting
it – is oriented towards synthesis, analytical reductionism being only a – however
important and indispensable – step in this direction. The concept of ecosystem is
capable of synthesising, or integrating, its main properties as revealed through its
analysis: dynamic connections, transience, scale-dependent views, irreversibility,
short-term predictability and long-term unpredictability. The concept will therefore
be an indispensable tool for the – hopefully sustainable – development of human-
kind in the twenty-first century.
References
Allen TFH, Hoekstra TW (1990) The confusion between scale-defined levels and conventional
levels of organization in ecology. J Veg Sci 1:5–12
Allen TFH, Zellmer AJ, Wuennenberg CJ (2005) The loss of narrative. In: Cuddington K, Beisner
BE (eds) Ecological paradigms lost. Routes of theory change. Elsevier Academic Press,
Burlington, pp 333–370
Aristotle (1995) Poetics, ed. and transl. by S Halliwell. Harvard University Press, Cambridge
Mass
Bergandi D (1995) “Reductionist holism”: An oxymoron or a philosophical chimera of E. P. Odum’s
systems ecology? – Ludus Vitalis, Revista de filosofia de las ciencias de la vida.
J Philos Life Sci 3(5):145–180, Mexico /Barcelona
Blandin P (2006) L‘écosystème existe-t-il? Le tout et la partie en écologie. In: Gayon J, Martin T
(eds) Le tout et la partie. CNRS Editions, Paris
Blandin P, Lamotte M (1988) Recherche d’une entité écologique correspondant à l’étude des
paysages: la notion d’écocomplexe. Bulletin d’écologie 19:547–555
Breckling B, Müller F (1997) Der Ökosystembegriff aus heutiger Sicht - Grundstrukturen und
Grundfunktionen von Ökosystemen. In: Fränzle O, Müller F, Schröder W (eds) Handbuch der
Umweltwissenschaften. Ecomed Verlagsgesellschaft, Landsberg, chapter II-2.2
Brussard PF, Michael Reed J, Richard Tracy C (1998) Ecosystem management: what is it really?
Landsc Urban Plann 40:9–20
Cantlon JE (2002) Ecological bridges revisited. Bull Ecol Soc Am 83:271–272
Carpenter RA (1995) A consensus among ecologists for ecosystem management. Bull Ecol Soc
Am 76:161–162
Cherrett JM (1989) Key concepts: the result of a survey of our members’ opinions. In: Cherrett
JM (ed) Ecological concepts. The contribution of ecology to an understanding of the natural
world. Blackwell, Oxford, pp 1–16
226
W. Haber
Daily GC (1997) Nature’s services: Societal dependence on natural ecosystems. Island Press,
Washington, DC
De Laplante K (2005) Is ecosystem science a postmodern science? In: Cuddington K, Beisner BE
(eds) Ecological paradigms lost. Routes of theory change. Elsevier Academic Press,
Burlington, pp 397–416
Egler FE (1942) Vegetation as an object of study. Philos Sci 9:245–260
Ellenberg H (1973) Versuch einer Klassifikation der Ökosysteme nach funktionellen
Gesichtspunkten. In: Ellenberg H (ed) Ökosystemforschung. Teil VII, Die Ökosysteme der
Erde. Springer, Berlin, pp 235–265
Ellenberg H, Mayer R, Schauermann J (eds) (1986) Ökosystemforschung. Ergebnisse des Solling-
Projektes 1966–1986. Ulmer, Stuttgart
Evans FC (1956) Ecosystem as the basic unit in ecology. Science 123:1127–1128
Evans GC (1976) A sack of uncut diamonds: the study of ecosystems and the future resources of
mankind. J Ecol 64:1–39
Feibleman JK (1954) Theory of integrative levels. Br J Philos Sci 5:59–66
Ford ED, Ishii H (2001) The method of synthesis in ecology. Oikos 93:153–160
Fränzle O (1998) Grundlagen und Entwicklung der Ökosystemforschung. In: Fränzle O, Müller
F, Schröder W (eds) Handbuch der Umweltwissenschaften, Part 3–2.1. Ecomed, Landsberg,
pp 1–24
Fränzle O (2001) Alexander von Humboldt’s holistic world view and modern inter- and transdis-
ciplinary ecological research. Northeast Nat 1:57–90, Special Issue
Golley FB (1993) A history of the ecosystem concept in ecology. More than the sum of the parts.
Yale University Press, New Haven
Haber W (2004) Landscape ecology as a bridge from ecosystems to human ecology. Ecol Res
19:99–106
Hölter F (ed) (2002) Scales, hierarchies and emergent properties in ecological models. Peter Lang,
Berlin
Holling CS (1992, 1996) Cross-scale morphology, geometry, and dynamics of ecosystems. Ecol
Monogr 62:447–502, and In: Samson, F.B. and F.L. Knopf 1996. Ecosystem management.
Selected readings. Springer, New York, pp. 351–423
Jax K (2002) Die Einheiten der Ökologie. Peter Lang, Frankfurt/M
Johnson NC, Malk AJ, Sexton WJ, Szaro RC (1999) Ecological stewardship. A common reference
for ecosystem management. Elsevier, New York
Likens GE (1992) The ecosystem approach: its use and abuse. Ecology Institute, Oldendorff/
Luhe
Lindeman RL (1942) The trophic-dynamic aspect of ecology. Ecology 23:399–418
Malone CR (1995) Ecosystem management: Status of the federal initiative. Bull Ecol Soc Am
76:158–161
Mayr E (1997) This is biology. The science of the living world. Belknap Press of Harvard
University Press, Cambridge
MEA (Millennium Ecosystem Assessment) (2003) Ecosystems and human well-being. Island
Press, Washington, DC
MEA (Millennium Ecosystem Assessment) (2005) Ecosystems and human well-being: Synthesis.
Island Press, Washington, DC
Mueller-Dombois D, Ellenberg H (2002) (Reprint 1974): Aims and methods of vegetation ecol-
ogy. Blackburn Press, Caldwell
Novikoff AB (1945) The concept of integrative levels and biology. Science 101:209–215
Odum EP (1953) Fundamentals of ecology. First edition. 1959: Second edition. 1972: Third edi-
tion. Saunders, Philadelphia
O’Neill RV (2001) Is it time to bury the ecosystem concept? (With full military honors, of
course!). Ecology 82:3275–3284
O’Neill RV, DeAngelis DL, Waide JB, Allen TFH (1986) A hierarchical concept of ecosystems.
Princeton University Press, Princeton
227
Peterson GD (2005) Ecological management: Control, uncertainty, and understanding. In:
Cuddington K, Beisner B (eds) Ecological paradigms lost. Routes of theory change. Elsevier
Academic Press, Burlington, pp 371–395
Picht G (1979) Ist Humanökologie möglich? In: Eisenbart C, Eisenbart C (eds) Humanökologie
und Frieden. Klett-Cotta, Stuttgart, pp 14–123
Pomeroy LR, Alberts JJ (eds) (1988) Concepts of ecosystem ecology. A comparative view.
Springer, New York
Potthast T (2002) From “mental isolates” to “self-regulation” and back: justifying and discovering
the nature of ecosystems. In: Schickore J, Steinle F (eds) Revisiting discovery and justification.
Preprint 211. Max Planck Institute for the History of Science, Berlin, pp 129–142
Reiners WH (1986) Complementary models for ecosystems. Am Nat 127:59–73
Samson FB, Knopf FL (1996) Ecosystem management. Selected readings. Springer, New York
Schaffer WM (1985) Order and chaos in ecological systems. Ecology 66:93–106
Scheiner SM, Hudson AJ, van der Meulen MA (1993) An epistemology for ecology. Bull Ecol
Soc of Am 74:17–21
Schultz AM (1967) The ecosystem as a conceptual tool in the management of natural resources.
In: Ciriacy-Wantrup SV, Parsons JJ (eds) Natural resources, quality and quantity. The
University of California Press, Berkeley, pp 139–161
Schwarz AE (2003) Wasserwüste, Mikrokosmos, Ökosystem. Rombach, Freiburg
Sheail J (2005) Tansley and British ecology: The formative years. Bull Br Ecol Soc 36:23–25
Sheifer IC (1996) Integrating the human dimension in ecosystem/ecoregion studies – a view from
the ecosystem management national assessment effort. Bull Ecol Soc Am 77:177–180
Stein SM, Gelburd D (1998) Healthy ecosystems and sustainable economies: the federal inter-
agency ecosystem management initiative. Landsc Urban Plann 40:73–80
Szaro RC, Sexton WT, Malone CM (1998) The emergence of ecosystem management as a tool for
meeting people’s needs and sustaining ecosystems. Landsc Urban Plann 40:1–7
Tansley AG (1922) Elements of plant biology. George Allen & Unwin, London
Tansley AG (1935) The use and abuse of vegetational concepts and terms. Ecology 16:284–307
Taylor P, Haila Y (2001) Situatedness and problematic boundaries: conceptualizing life’s complex
ecological context. Biol Philos 16:521–532
Trepl L (1988) Gibt es Ökosysteme? Landschaft Stadt 20:176–185
Vogt KA, Gordon JC, Wargo JP, Vogt DJ, Asbjornsen H, Palmiotto PA, Clark HJ, O’Hara JL,
Keaton WS, Patel-Weynard T, Witten E (1996) Ecosystems. Balancing science with manage-
ment. Springer, New York
Weil A (2005) Das Modell “Organismus” in der Ökologie: Möglichkeiten und Grenzen der
Beschreibung synökologischer Einheiten, vol 11, Theorie in der Ökologie. Peter Lang,
Frankfurt/M
Weinberg GM (1975) Introduction to general systems thinking. Wiley, New York
Wiegleb G (1996) Konzepte der Hierarchie-Theorie in der Ökologie. In: Mathes K, Breckling B,
Ekschmitt K (eds) Systemtheorie in der Ökologie. Ecomed, Landsberg, pp 7–25
Wilcove DS and F B Samson (1987) Innovative wildlife management: listening to Leopold. Trans
North Am Wild Nat Resour Conf 52: 321–329
Willis AJ (1997) Ecology of dunes, salt marsh, and shingle. Chapman & Hall, London and
New York
Wilson EO (1996) In search of nature. Island Press, Washington, DC
Zeide B (1996) Is “The scientific basis” of ecosystem management indeed scientific? Bull Ecol
Soc of Am 77:123–124
18 An Ecosystem View into the Twenty-first Century
Part VI
Local Conditions of Early Ecology
231
Chapter 19
Early Ecology in the German-Speaking
World Through WWII
Astrid Schwarz and Kurt Jax
The scientific practice and theory of ecology in the German-speaking world arose
simultaneously yet independently of each other in different places and in relation
to different subjects. The new disciplining perspective took in lakes and fish ponds
as well as native forest, heath and mountain landscapes, though it also included the
flora and fauna of tropical and arctic regions. “German-speaking world” refers here
not so much to an area determined by its political or natural borders but rather by
its linguistic boundaries. A lively exchange of publications, objects and individuals
took place within this scientific world. Cities and regions belonging to different
spheres of political influence were a part of this Sprachraum, which encompassed
Zurich, Vienna, Prague, Budapest and Berlin, as well as Bohemia, Silesia and
Prussia, the Rhineland and the Valais. Perfect examples of the commonplace
exchanges that took place in what we call the “German-speaking world” of that
time were the botanists Simon Schwendener and Gottlieb Haberlandt, who were
decisive for the formation of physiological plant ecology (see below). Schwendener
was born and educated in Switzerland and spent most of his working life in
Germany (Tübingen and Berlin; prior to that in Basel, Switzerland); Haberlandt
was born in Hungary, educated in Austria and worked for most of his life in Austria
(Vienna and Graz), though at times also in Germany (Tübingen and Berlin). So if
– for the sake of brevity – we speak of “German” ecology in this chapter, we mean
this region as delimited by the common use of the German language as a means of
communication.1
1 Even today there is a joint Society of German-speaking ecologists, the Gesellschaft für Ökologie
(GfÖ), founded in 1978, which includes scientists mainly from Germany, Austria, Switzerland
and Liechtenstein.
A. Schwarz (*)
Institute of Philosophy, Technische Universität Darmstadt, Schloss, 64283 Darmstadt,
Germany
e-mail: schwarz@phil.tu-darmstadt.de
K. Jax (*)
Department of Conservation Biology, Helmholtz Centre for Environmental Research (UFZ),
Permoserst.15, 04318 Leipzig, Germany
e-mail: kurt.jax@ufz.de
A. Schwarz and K. Jax (eds.), Ecology Revisited: Reflecting on Concepts, Advancing Science,
DOI 10.1007/978-90-481-9744-6_19, © Springer Science+Business Media B.V. 2011
232
A. Schwarz and K. Jax
Indeed up until the first 20–30 years of the twentieth century, German was
still one of the dominant languages of scholarly research, including the natural
sciences. The international symposium “Deutsch als Wissenschaftssprache im
20. Jahrhundert” (German as a language of scholarship in the twentieth century),
held in January 2000 and organized by the Mainz Academy of Sciences and
Literature, arrived at the conclusion that “[f]rom the mid-nineteenth century
through to the 1920s German was a world language for the sciences; after this –
from the 1930s due to the Nazi policy of expulsion and from the late 1950s due
to international developments – it became a marginal language, like French and
Russian”.2 Thus in the period up until the Second World War there were scien-
tific journals containing for the most part articles written in German – including
pieces by non-native speakers. However, one would also come across isolated
articles written in English or French in these journals, indicating that they were
widely recognized publication forums for scientists throughout the world.
Among these, for example, is the Biologisches Centralblatt (founded in 1881),
published since 1997 under the title Theory in Biosciences, Engler’s Botanische
Jahrbücher (founded in 1880) and the Zoologische Anzeiger (founded in 1878)
but also more popularly oriented journals such as the Kosmos founded in 1877
by Ernst Haeckel to disseminate the Darwinian Weltanschauung. These journals
were part of what was called the “second wave of formation”3 of natural history
journals and they were all important forums for an evolving German-language
ecology. In addition to these journals, the many varied publications of the “sci-
entific societies” were also used to disseminate ecological ideas and scientific
programmes.4
Just a short time later, the first journals to specialize in ecological topics were
founded. The spectrum of journals diversified rapidly, encompassing terrestrial and
aquatic ecology and reflecting the growing consolidation of the new field of
research. The founding of more journals at the start of the twentieth century con-
tributed towards the increasing stabilization of the field and a clearer delineation of
ecological research questions. The first specialist ecological journals included the
Forschungsberichte der Biologischen Station zu Plön, which continued to exist
from 1906 onwards under the title Archiv für Hydrobiologie und Planktonkunde; in
2 “Pörksen 2001, p. 29.”
3Andreas W. Daum speaks of a “zweite Gründungswelle”of natural history journals in 19th century,
especially in the domain of scientific popularization (Daum 1998, p. 359).
4 There are numerous examples of these in the 1880s and 1890s especially, but exactly how publi-
cation strategy related to the constitution of the community – particularly with regard to the
relationship between “lay people”, independent scholars and employed scientists – has not yet
been systematically studied. Those societies active in the publishing sphere include the scientific
societies (Naturforschende Gesellschaften) in Zurich, Lucerne and Lausanne as well as the
Natural History Society of the Prussian Rhineland and Westphalia (Bonn). A more general
perspective on the popularization of science is given for instance in Daum (1998), who deals in
one chapter also with the “Naturvereine” in 18th century.
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19 Early Ecology in the German-Speaking World Through WWII
1920 it was re-named the Archiv für Hydrobiologie5; the first volume of the
Internationale Revue der gesamten Hydrobiologie und Hydrographie was pub-
lished in 1908, while from 1923 onwards the International Association of
Theoretical and Applied Limnology (IVL) produced a Zeitschrift der Internationalen
Vereinigung für theoretische und angewandte Limnologie; the Geobotanic Institute
Rübel in Zurich also had its own journal in 1924, as did the Biological Institute
Helgoland, with its Helgoländer wissenschaftliche Meeresuntersuchungen, from
1937 onwards.
Shaping the Field
Thus the first steps toward a scientific ecology in the German-speaking world can
be dated back to the second half of the nineteenth century. They were taken by bota-
nists and zoologists, microbiologists and physiologists, hydrologists, geographers
and chemists. From about the 1870s onwards an increasing number of research
programmes were devised and scientific networks created around people such as
zoologist Karl Möbius (1825–1908) and physiologist Victor Hensen (1835–1924)
at the University of Kiel, Anton Frič (1832–1913) at the Institute of Prague
University, botanist Carl Schröter (1855–1939) at the University of Zurich,
Friedrich Zschokke (1860–1936) at the Institute for Zoology in Basel, and within
the study group around botanist Simon Schwendener (1829–1919) in Berlin. The
formation of networks was further supported by the founding of specialist commis-
sions, whether it occurred in scientific societies, in organizations, or through politi-
cal interventions.
For example, in 1870 the Prussian government appointed a commission for the
scientific exploration of the German oceans (Kommission zur wissenschaftlichen
Erforschung der deutschen Meere), of which not only Hensen and Möbius were
members but also plant geographer Adolf Engler, anatomist Carl von Kupffer,
physicist Gustav Karsten and other scientists. A “limnological commission” was
set up at a meeting of the Schweizerische naturforschende Gesellschaft (Swiss
Scientific Society) in Frauenfeld in the year 1887, whose task was to develop, coor-
dinate and organize specific research projects and to procure and archive as well as
5 This step-by-step process of re-naming can be seen as a means of asserting a continuity in editor-
ship: Volume 10 from 1915 was the last to be edited by Otto Zacharias on his own; Volume 11
(1917) begins with an obituary for Zacharias written by August Thienemann, the new co-editor;
and both are named in Volume 12 from 1920, the point at which the journal changed its name to
Archiv für Hydrobiologie. Not until Volume 13 (1922) does August Thienemann appear as the
sole editor; this is also the year in which the International Association of Theoretical and Applied
Limnology was founded in Kiel. August Thienemann was the co-initiator, along with Einar
Naumann (the “junior partner”), of this society.
234
A. Schwarz and K. Jax
6 The initiator was François Auguste Forel. While in 1890 the commission still comprised three
official members, in 1892 it had grown to twice this size. In addition to Forel, its members were:
Forests Inspector Coaz from Bern (from 1887), Prof. F. Zschokke from Basel, Dr. E. Sarasin and
Prof. L. Duparc from Geneva, and grammar school teacher Prof. X. Arnet from Lucerne. The
composition of the commission is representative of early ecological research in two respects: first,
it is interdisciplinary and, second, its members come from both academic and non-academic
milieus (Proceedings of the Swiss Scientific Society, 74th Annual Meeting in Freiburg, Annual
Report 1890–1891, Freiburg: Gebr. Fragnière 1892, pp. 100–103, 112–115, 142).
7 Halbfass’ negotiations with the representatives of the German Geographers Congress and the
re-shaping of the text to become a publishable, politically correct version are recounted in Ein
vergessenes Kapitel aus der Seenforschung (A forgotten chapter in the history of lake research)
(2005) by Sylvin Müller-Navarra, pp. 188ff.
8 An insightful discussion about the role of German-based Natural History Museums is given in a
comparative study by Carsten Kretschmann (2006) Räume öffnen sich (spaces open up).
9 “Die aus Liebe zur Wissenschaft und aus Patriotismus arbeitenden Kräfte auf einen kargen Ersatz
der Barauslagen angewiesen. […] in Deutschland (stehen) dem Unternehmen Tausende zur
Disposition.” Interestingly enough, Frič continues here: “Repeated attempts to encourage the large
landowners to support this endeavour for the common benefit have borne no fruit” (Wiederholte
Versuche, den Grossgrundbesitz zur Förderung dieses gemeinnützigen Vorgehens aufzumuntern,
führten zu keinem Resultat) (queryFrič and Václav Vávra 1894, p. 7). Apparently, however, there
were exceptions, as Frič also reports that Freiherr von Dercsényi had a “solidly built nice little
house” (festgebautes, nettes Häuschen) erected, which then served as a zoological research station.
10 The research results from these activities were published in the Archiv für die naturwissenschaftli-
che Landesdurchforschung von Böhmen. The latter also contained, in 1894, Anton Frič’s
Untersuchungen über die Fauna der Gewässer Böhmens, which essentially referred to two artificial
bodies of water, the Unterpocernitz and the Gatterschlag Pond. However, the study of Bohemian
forest lakes and other artificial bodies of water had been initiated much earlier, in 1872 (Frič and
Václav Vávra 1894, pp. 5–7).
facilitate publications.6 Wilhelm Halbfass (1856–1938) gave a talk at the 13th
German Geographers Congress in 1901 in Breslau entitled “The scientific and
economic significance of regional limnological institutes” (Die wissenschaftliche
und wirtschaftliche Bedeutung limnologischer Landesanstalten). The talk ended
with a “resolution”, addressed to the Prussian government, calling for the establishment
of regional limnological institutes. This call for action, its political content toned
down, was given a forum in Petermanns Mitteilungen in 1902: it was now limited
to an exhortation to gather data on all the lakes in Europe.7
Likewise dedicated to “naturwissenschaftliche Landesdurchforschung” (regional
scientific research) were comités set up in Austria-Hungary (their headquarters in
Vienna and Prague), which were concerned mainly with cartographical images but
also with determining national resources, including not only geological formations
but also “the animal world of our lakes, ponds and rivers”. Zoologist Anton Frič,
secretary of the Bohemian initiative that emerged from the Prague-based museum
milieu,8 lamented nonetheless that “those working out of love for science and out
of patriotism have to rely on a meager reimbursement of their cash expenses” while
“in Germany the enterprise has thousands at its disposal”,9 naming in this context
the “zoological station at Ploen”.10 This Biologische Station zu Plön, established in
1892, was one of the nodal points of activity around which the new field of research
became institutionalized. Other early initiatives included the founding of the
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19 Early Ecology in the German-Speaking World Through WWII
Biologische und Fischerei-Versuchsstation Müggelsee (Biological and Fisheries
Experimental Station at Müggelsee) in Berlin-Friedrichshagen (1893) and the
Biological Station in Lunz, Austria in 1906: all these contributed towards the insti-
tutionalization of ecological research.
“Limnology” – An Attractor in the Emerging Field?
The coming together of ecologists in the International Association of Theoretical
and Applied Limnology (IVL) was a further step in establishing aquatic ecology11 as
a scientific discipline as well as in the perceptions of the wider society. In 1922, the
first congress took place in Kiel; it was conducted and also recorded in writing in
German – “as is customary for international congresses – in the language in which
they are held”, writes Secretary General Friedrich Lenz (1889–1972) in the
Foreword, which is also printed as a Preface and an Avant-Propos. The participants
at the inaugural conference numbered 67 members and 15 guests from a total of
13 countries. In 1923, however, when the Proceedings were published, more than
350 “full members” from 26 countries were registered,12 along with 31 institutions
as “extraordinary members”, as can be seen from the members directory in this first
volume of the “Proceedings from the International Association for Theoretical and
Applied Limnology”. The themes addressed include the metabolic rate in open
water and near the bank – the so-called pelagic and littoral zones – and the role of
bacteria and funghi in these processes. The metabolic characterization, or circadian
rhythm, of plankton and its origins are discussed along with the problems of the
fisheries and of hydraulic engineering, water quality and the protection of lakes and
rivers. The newly founded society could count on a certain level of support in the
public perception when it came to the protection of aquatic sites in particular, backed
up at an institutional level by the Heimatschutz- and Naturschutzvereine roughly
translated, the local societies for the protection of the country and conservation of
nature. However, local water supply and wastewater disposal companies were also
affected by the problems attending water pollution and water usage – as, indeed,
were all those responsible for the administration of water as a resource at the
national governmental level. The production of drinking water and the use of hydro-
power to generate electricity had already long become a matter of national interest
in the nineteenth century, which made them a potential source of conflict. Indeed
disputes were rife between Heimatschutz, state authorities and industry. Plans to
11 Aquatic ecology here describes above all the study of inland waters, although marine biologists
were also involved in establishing the IVL (August Pütter and Ernst Hentschel as well as Karl Brandt
from the Oceanographic Institute at Kiel attended the 1922 conference in Kiel). Marine biology had
been established prior to limnology, which initially disassociated itself from the former for reasons
that had more to do with competition for research funding and prestige; it also maintained that it had
a more progressive research programme: “…la limnologie peut appliquer l’expérimentation là où
l’océanographie en est le plus souvent réduit à la seule observation.” (Forel 1896, p. 596).
12 In his opening speech Thienemann speaks of another 187 individuals who had responded to the
mailing of 1,000 copies of “Calls to join” – a response to a campaign initiated by Naumann and
Thienemann which was regarded as highly positive (1923, p. 1).
236
A. Schwarz and K. Jax
build the barrage on the Rhine at Lauffenburg (1914) using dynamite and dam con-
structions were contested, but in the end the barrage did get built. On the other hand,
in 1942 another disputed project, the so-called Rheinwald three-stage project on the
posterior Rhine as well as the building of a power station were stopped after a long
dispute (Tümmers 1999, pp. 38 f., pp. 88 f.). This ongoing process of industrializa-
tion along the rivers also left behind traces that could not only be seen but also smelt.
In 1901 the issue of the Rhine being a “cesspool” was debated in the German
Reichstag. The conflicts over the quality and production of drinking water quickly
led to the establishment of state institutions. The Royal Experimentation and Testing
Institute for Water Supply and Wastewater Removal (Königliche Versuchs- und
Prüfunganstalt für Wasserversorgung und Abwasserbeseitigung) was founded in
Berlin in 1901 (Kluge and Schramm 1986, pp. 84 ff.).
One of the co-founders of the IVL, August Thienemann (1882–1960), was quick to
recognize this interrelationship between politics and science and between industry and
the state in the sphere of ecological research; he sought to make use of it not only in
conceptual terms but also as a tool for institution building. Thienemann wanted limnol-
ogy to be seen as a “bridging science” (Brückenwissenschaft), for herein, according to
him, lay “its great cultural significance for our times” (1935, p. 20). Thienemann bor-
rows this term from a piece entitled Die Struktur der Ganzheiten (The Structure of
Wholes) by philosopher Wilhelm Burkamp (1929), a text from which he frequently
quotes (for instance 1933; 1935). What seems especially noteworthy to Thienemann is
Burkamp’s characterization of new sciences as problem-based, as “methodological
and factually structural wholes” (methodische und sachlich strukturelle Ganzheiten”,
Burkamp quote, emphasis AS, KJ), as this supports his own formulation in which the
unique character of limnology is seen to be grounded in “the [research] object and in
the methods” (1935, p. 19; emphasis in original).
With this statement, Thienemann was effectively placing limnology explicitly in
a mediating position between various natural science disciplines, predominantly
hydrography, biology, geology and oceanology, as well as between basic and applied
research. “Theory will always remain the foundation for practice!” he exclaims, adding
as a caveat that nonetheless “the study of wastewater (an extreme milieu with regard
to its chemistry), which was originally undertaken exclusively for practical reasons,
has made it quite considerably easier for us to gain a theoretical understanding of the
population in a chemically normal natural bodies of water” (1923, p. 3).13 What is
13 “…das Studium der Abwässer, eines in chemischer Hinsicht ganz extrem gestalteten Milieus,
das ursprünglich doch ausschließlich unter praktischen Gesichtspunkten vorgenommen wurde,
uns das theoretische Verständnis der Besiedelung chemisch-normaler natürlicher Gewässer ganz
wesentlich erleichtert hat”; Thienemann’s own research work was located initially in both the
theoretical and the applied sphere. Later, however, and certainly after his definitive decision to
work at the Kaiser-Whilhelm-Institut (KWI) Plön – the former Biologische Station zu Plön - rather
than to succeed Bruno Hofer at the Royal Bavarian Biological Testing Station for Fisheries
(Königliche Bayerische Biologische Versuchsanstalt für Fischerei) in Munich, his research can be
described more as theoretical in the sense of conceptual work. Nonetheless he remained active in
fisheries biology, in committees and organizations, and reference was frequently made to his work
on biological studies of wastewater. In 1956 the Department of Agriculture and Horticulture at
Humboldt University, Berlin conferred on him the degree of Dr. agrar. h.c.
237
19 Early Ecology in the German-Speaking World Through WWII
especially remarkable about this statement is that, rather than supporting the familiar
idea of knowledge transfer from theory to practice, or from the natural sciences to
the engineering sciences, it also recognizes the reverse trajectory and acknowledges
it as a source of cognitive insight and as a fruitful heuristic.
Thus Thienemann offers not only methodological arguments for the bridging
function of limnology or justifies it by reference to a specific set of research objects;
he also argues that the conceptual framework of this new science enables it to function
as a bridge. He brings into play the link to concepts of natural philosophy, taking up once
again a romantic conception of nature; at the same time, though, he also mentions
modernity and the progressiveness and experimental character of limnology, which he
claims makes it different from the “merely” descriptive sciences of natural history.
Thienemann returns again later to the term “bridging science” in order to char-
acterize ecology, whose purpose, as the study of nature’s household, is to connect
“all the branches of the study of nature” (“alle Zweige der Naturkunde”).14 In the
same year he also uses the metaphor Grenzland Limnologie (borderland limnology) – a
place located between the “motherland” of biology and physiography. Much like
the term “bridging science”, the notion of a “borderland”, or border zone, or even
“border science” (Grenzwissenschaft) are part of a stabilizing terminology which
serves to describe ecology to this day.15
In his attempts to describe ecology in conceptual terms, Thienemann refers
repeatedly to limnology. This is the case, for example, with his use of the conceptual
scheme of the three stages of ecology, which he apparently published for the first
time in his 1942 article Vom Wesen der Ökologie (On the essence of ecology). This,
too, is an attempt to give ecology a conceptual framework (given that it was a field
still relatively untried in scientific practice), using limnology as a model example of
an integrating science; this he could do because limnology was already institutionalized
and possessed recognizable contours in the form of research programmes (Fig. 19.1).
14 Thienemann 1942, p. 324. In his obituary, G.H. Schwabe stresses that Thienemann’s legacy lay
in seeing the core task of ecology as one of nurturing links – not only as a bridging science
between scientific disciplines, but also between the natural sciences and humanities, between
small and large interdisciplinarity, so to speak. Schwabe himself – treading to an extent in
Thienemann’s footsteps – situates himself in the tradition of a holistic world view and sets his
sights on founding a meaning-making ecology, which he believes capable of preventing “modern
civilization from drifting without an anchor into the incomparable”. He continues: “[E]cology is
a logically necessary connecting link between the natural sciences and the humanities and as such
is inescapably at the mercy of the tensions and conflicts that are in keeping with its very essence”
([D]ie Ökologie [ist] ein logisch notwendiges Bindeglied zwischen Natur- und Geisteswissenschaften
und als solches unvermeidlich den Spannungen und Konflikten ausgesetzt, die seinem Wesen
entsprechen; 1961, p. 316).
15 Thienemann 1927, p. 33. Historian Robert Kohler has developed the concept of “border zone”
ecology where objects, concepts and individuals constantly travel back and forth between lab and
field (Kohler 2002). The bridge metaphor, too, is taken up time and again as a means of character-
izing ecology. The tradition and tenability of the semantics of bridging are reconstructed by Hans
Werner Ingensiep and Thomas Potthast in “Brückenschläge – zur Sprache der Ökologie” (Building
bridges: On the language of ecology) and “Ökologie’ als Brücke zwischen Wissen und Moral?”
(‘Ecology’ as a bridge between knowledge and morality?) (Busch (ed.) 2007).
A. Physiographischer Teil
Die chemischen und
physikalischen Eigenschaften
des Wassers
Das Einzelleben
im Wasser
B. Biologischer Teil
Das Gemeinschaftsleben in
den Binnengewässern
Das Gesamtleben der Binnen-
gewässer
(Biotop und Biocoenose in
Wechselwirkung und als
Einheit.)
Die hydrographischen und
geographisch-geologischen
Eigenschaften der limnischen
Biotope
II. Coenographische Stufe
III. Limnologische
Stufe
I. Idiographische Stufe
Die drei Stufen der limnologischen Forschung.
a
Das Gesamtleben der Binnengewässer
(Biotop und Biocönose in
Wechselwirkung und als
Einheit)
(Lebenseinheit: die Biocönose)
Das Gemeinschaftsleben in
den Binnengewässern
Die hydrographischen und
hydrogeographischen
Eigenschaften der Gewässer
Die chemischen und
physikalischen Eigenschaften
des Wassers
(Lebenseinheit: die Art)
Das Einzelleben
im Wasser
B. Biologischer Teil A. Physiographischer Teil
III. Limnologische stufe
II. Cönographische Stufe
I. Idiographische Stufe
Die drei Stufen der limnologischen Forschung.
b
Fig. 19.1 (continued)
239
19 Early Ecology in the German-Speaking World Through WWII
Allgemeine Ökologie
oder die
Lehre vom Haushalt der Natur
Lebenseinheit:
Lebenseinheit:
Lebenseinheit:
Die lebenerfüllte Lebensstätte
Die Organismenart
Die Lebensgemeinschaft
oder Biocönose
Biocönotik
oder die
Lehre vom Gemeinschaftsleben Die Standortsbedingungen
der einzelnen Biotope
II. Conographische Stufe
III. Holographische Stufe
I. Idiographische Stufe
Autökologie
oder die
Lehre vom Einzelleben
Die Eigenschaften
von Wasser und Luft
als Lebensbedingungen
B. Biologischer Teil A. Physiographischer Teil
Die drei Stufen der Ökologie.
c
Fig. 19.1 (continued) (a) “The three stages of limnological research” (Thienemann 1925, p. 680)
(b) “The three stages of limnological research” (Thienemann 1935, p. 18) are placed upside down
and reversed sideways and are thus corrected, following the disciplinary mode of reading: In lim-
nological research B goes ahead of A – the biological part takes precedence over the physiographic
part (c) “The three stages of ecology” (Thienemann 1942, p. 325). In the text, autoecology is iden-
tified with the idiographic stage and synecology with the coenographic stage, although Thienemann
agrees with Friederichs when he states: “Autoecology is a sub-concept of synecology”, seeing in
this a confirmation of his hierarchical three stages of integration (Thienemann 1942, p. 316)
Thienemann was careful to anchor the programmatic element of “his limnology”
within a bourgeois educational canon, which included as a matter of course a Goethe-
inspired natural philosophy and a set of Christian motifs.16 Typically an encounter
occurs here between cosmological and romantic – even sacred – elements. There is
16The emphasis placed on the personality of the researcher as the key ingredient which gives
ecological knowledge that certain something to make it especially valuable and thus to set it apart
from “just” making the scientific claim to objectivity was also a part of the bourgeois self-image:
“Every scientific study that rises above the average has a markedly personal, that is to say, a
subjective touch; and it is often this that makes such a study especially interesting. This is why I
think we can only really fully appreciate the work of our scientific colleagues when we know them
personally” (Jede wissenschaftliche Arbeit, die sich über das Durchschnittsniveau erhebt, hat eine
starke persönliche, also subjektive Note, und diese ist es oft, die eine solche Arbeit besonders
interessant macht. Deshalb meine ich, können wir die Arbeiten unserer Fachgenossen erst voll
werten, wenn wir unsere Fachgenossen persönlich kennen; Thienemann 1923, p. 4 f.).
240
A. Schwarz and K. Jax
much talk, for example, of the “contemplation of the small with a view to the large”
(Andacht zum Kleinen mit dem Blick aufs Große), that “the whole always exists prior
to the parts” (das Ganze stets vor den Teilen da ist) and that “the parts are a world unto
themselves, and yet intermesh with one another harmoniously” (die Teile jeweils eine
Welt für sich sind, aber doch harmonisch ineinander greifen), while the lake is
described as an “arena of life” (Bühne des Lebens) or as “a world in miniature” (eine
Welt im Kleinen). Thienemann, for example, repeatedly establishes the connection to
Goethe – this too occurring within a decades-old continuity that outlasts all political
and epistemic breaks: In 1939 the “primary axiom of the holistic world view” (erste
Axiom ganzheitlicher Weltauffassung; Meyer-Abich 1938 in a preface to Smut’s Die
holistische Welt) is “verified” by means of a quote from Goethe: “How everything
weaves itself into the whole, one in the other works and lives” (Wie alles sich zum
Ganzen webt, Eins in dem andern wirkt und webt; 1939, p. 12). This very sentence is
also quoted in 1951, again with the reference to the “axiom of holism”; indeed the
“intellectual world of Goethe also lives on in today’s natural science, according to the
authoritative judgment of modern scientific researchers”17 (1951, p. 580). Finally, to
name one last example, in 1954 we find this: “Our intellectual attitude towards nature
must not be exhausted in the endeavour to identify its laws as the foundation of our
material culture; instead, more than this, it must strive upwards to encompass a view
of nature such as the one into which Goethe breathed life, who saw in living nature and
in every one of its parts the whole, the great harmony in all disharmonious separate
phenomena, and to whom it was a well-ordered whole, a cosmos”18 (1954, p. 49).
This is the language of many, if not the majority, of the scientists in the formative
ecological community. It is a style of writing distinguished by a remarkable degree of
continuity and can still be found before, during and even after World War II. The titles
of a few selected publications, again by August Thienemann, may serve to illustrate
this: 1937 Lebensgemeinschaft und Lebensraum (Living community and living
space), 1939 Grundzüge einer allgemeinen Ökologie (Principles of a universal ecology)
accompanied by the motto “Gemeinschaft ist die Lebensform der Natur” (Community
is nature’s way of life),19 1944 Der Mensch als Glied und Gestalter der Natur (Man
as part of and shaper of nature), 1951 Vom Gebrauch und vom Mißbrauch der
Gewässer in einem Kulturlande (Of the use and misuse of freshwater in a land of
culture), and 1954 Wasser: das Blut der Erde (Water: the blood of the earth).
17 “[die] geistige Welt Goethe’s [lebt] nach dem berufenen Urteil moderner Naturforscher auch fort
in der heutigen Naturwissenschaft”.
18 “Unsere geistige Haltung zur Natur darf sich nicht erschöpfen in dem Bestreben, ihre Gesetze
zu erkennen als Grundlage unserer materiellen Kultur; sie muß vielmehr darüber hinaus sich
emporringen zu einer Naturanschauung, wie sie Goethe beseelte, der in der lebenden Natur und
in jedem ihrer Teile stets das Ganze sah, die große Harmonie bei allen disharmonischen
Einzelerscheinungen, dem sie ein wohlgeordnetes Ganzes, ein Kosmos war”.
19 Originally an article in the Archiv für Hydrobiologie (1939, 35, pp. 267–285), the
Schweizerbart’sche Verlagsbuchhandlung publishing house also published a brochure bearing this
title. Of particular interest here is the form of the text which, apart from the closing comments, is
numbered in sections from 1–60, in similar fashion to a manifesto. Historian and philosopher of
241
19 Early Ecology in the German-Speaking World Through WWII
It is not hard to recognize that certain more or less radical nation-state tendencies
may also be implied here. For example, a passage from Thienemann’s writing states
“that the biological foundation for shaping and consolidating our German world
view must be the science, which explores the great interrelations in living nature,
namely: general ecology”20 (1942, p. 326). And almost 20 years earlier limnologist
Erich Wasmund (1902–1945) had written the following in an article on the scien-
tific provinces in the conservative journal Deutsche Rundschau (later instrumental-
ized by the National Socialist regime): “The language is a part of the style of the
innermost being of a people, and thus its specific scientific type becomes essential
to it. People and soil create a material conditionality of their scientific types”21
(1926, p. 245).
A further aspect of holistic conceptual figures becomes clear here, which serve
in some sense to elide the political dimension of the concept of nature. Nature is
treated as a moral category and is invested as such with an authority that cannot be
betrayed. Above all, though, whole and harmonious nature becomes an icon of a
way of thinking rooted in cultural pessimism which, during the Weimar period,
became an integral part of bourgeois self-perception and was further fuelled, for
example, by Oswald Spengler’s book The Decline of the West and by the philo-
sophical anthropology of Max Scheler, particularly his piece entitled Die Stellung
des Menschen im Kosmos (The place of man in the cosmos, 1928).
In this society, humans are perceived above all as destroyers of nature; at best they
assume the role of benevolent stewards, but nature can definitely not be a discursive
political term. This position – one that is critical of civilization and often technophobic
as a result – was also widespread in early ecology, where the more or less explicit
commitment to a holistic image of nature plays a central, integrating role and is more
or less a part of the philosophical core of the discipline. Yet a position such as this, sceptical
biology Thomas Potthast highlights the fact that this “little book” appeared “in greater detail” in
1941, entitled “Leben”, with several National Socialist slogans added in, while the new editions
published in 1956 and 1958 were cleansed linguistically of the National Socialist style “by [the
deletion of] about 10 passages” (2003, 252). What this proves is, first, the historical continuity of
the rhetoric of holism and its political malleability. Second, though, it also implies criticism of the
“mandarin” of ecology, Thienemann, expressed even more clearly at another point in the text:
“Currying favour with the powers that be, remaining silent on certain political practices, and
biased criticism are by no means necessarily mutually exclusive” (Anbiederung, Schweigen zu
bestimmten politischen Praktiken und partielle Kritik müssen sich […] keinesfalls ausschießen
[sic!]; 2003, 238). The volume Leben und Umwelt: Vom Gesamthaushalt der Natur (Life and
Environment: Of the total household of nature) was published in the series rowohlts deutsche
enzyklopädie (rde), which bore the motto “Paperback XXth Century Knowledge” and was designed
as such to achieve a broad public impact; the fact that it appeared in paperback form – quite a new
medium for the German reading public in the 1950s – was intended to contribute to this.
20 “daß für die Gestaltung und Festigung unserer deutschen Weltauffassung […] [die] biologische
Grundlage […] die Wissenschaft sein muß, die die großen Zusammenhänge in der lebenden Natur
ergründet: die allgemeine Ökologie.”
21 “Die Sprache ist aber Teil des Stils, Teil des Wesens eines Volkes, und so wird sein spezifischer
Wissenschaftstypus ihm wesentlich. […] Volk und Boden schaffen eine Materialbedingtheit ihrer
Wissenschaftstypen.”
242
A. Schwarz and K. Jax
towards civilization, is by no means necessary to a science geared towards holism,
either conceptually or politically.22 Accordingly, the understanding of holism contained
in the ecological body of knowledge gets played out in very varied ways: it may refer
more to practical knowledge, as envisaged by the planning sciences and scientific
nature conservation; or it may be elaborated in a more systems theoretical way, as with
the ecosystem models of 1950s American new ecology; or again it may present itself
as extremely technophilic, as with the large-scale experiment “Biosphere 2”.
Normalizing the Field: The Role of Textbooks
The first tentative steps taken in the new field of research were taken not within
academia alone but also, as mentioned above, beyond the confines of universities
and scientific institutions23: at high schools, museums, forestry, agriculture and fish
farming facilities, in the sphere of drinking water supply and that of medical
hygiene, as well as in the context of nature research societies in which lay people
and scientists engaged in lively exchange.24 In all these places, issues are addressed
which were later to be characterized as “ecological”. The natural science curricula
were not yet fixed either, so that quite a few ecologists who later became well-
known began their research initially as lay people; they included, for example,
Josias Braun-Blanquet and Wilhelm Halbfass, who went on to complete their scien-
tific career at a university or as internationally renowned independent scholars.
It is Ernst Haeckel who was credited with giving all these already existing activities
a label, even if he himself did not pursue any empirical work in the field which he
named “ecology” (see Chap. 16). The major contributions of German-speaking
researchers to the formation and maturing of early ecology came from other quarters,
before and after the publication of Haeckel’s “Oecologie” of 1866. The research projects
on the Baltic Sea and the North Sea, conducted by zoologist Karl August Möbius, are
a good example of this25; they are also a good example of the intertwining of theoretical
and applied activities in ecology, of scientific ecology and ecotechnology.
22 Joachim Radkau draws attention to the fact that in nature and environmental conservation cir-
cles during the 1930s and 1940s very varied positions regarding “nature” existed which are more
adequately described through reference to a polycratic model than to a theory of totalitarianism
(2003, p. 43).
23 Geographer and historian H.-G. Schultz makes the claim for geography that it was done mainly
outside scientific institutions in the first instance. In the case of ecology, this is true only for certain
areas, most obviously for those inspired by natural history and with their main focus on the observation
and description of isolated objects or events.
24 Unfortunately, this paper can not focus in detail on the relationship between academia and
learned societies, even if this is an important issue in early ecology (see, for instance, footnote 4).
The article by Patrick Matagne in this volume provides an initial sketch of this interrelationship,
whose significance has probably been underestimated.
25 Möbius’ Fauna of the Kieler Bucht, published in 1865, was a book that was seen by historians
of biology, as early as the 1920s, as furnishing ecology with a modern research programme and
methodology.
243
19 Early Ecology in the German-Speaking World Through WWII
Möbius began his field studies with a rather traditional programme of observation,
including classical zoological studies on a defined systematic group, but he also
addressed certain conceptual issues. This work was published in a book entitled
Fauna der Kieler Bucht in 1865.26 A short time later, in 1869, he was commissioned
by the Prussian government to study the oyster banks of Schleswig-Holstein; this was
related to the crisis in oyster mussel cultivation, where initial evidence suggested
overfishing had occurred. He developed a research programme, which eventually
resulted in the manuscript Die Auster und die Austernwirtschaft (The oyster and
oyster-culture) in 1877.27 It was this publication in which the term “biocoenosis” was
coined – not in the sense of a speculative blank concept but as a consequence of his
research experience and as a means to structure theoretical work in ecology as well.
Accordingly, in Chap. 10 – entitled An oyster bank is a biocoenosis, or living com-
munity (Eine Austernbank ist eine Biocönose oder Lebensgemeinde) Möbius writes:
Every oyster bank is in some sense a community of living creatures, a selection of species
and a sum of individuals which find in this very place all the conditions for their emergence
and maintenance, that is, the right soil, sufficient food, the appropriate salt content and
tolerable temperatures favourable for development.28
Thus the “oyster project” represented one of the first ecological programmes to
involve actual practical research, simultaneously influencing so-called applied sci-
ences such as fisheries and its political and economic environment. In addition,
however, biocoenosis represented a novel concept and method, which was taken up
by a wider public beyond academia, mainly in tertiary and secondary education.
A couple of books were published which navigated the boundary between textbook
and specialized work, between general and expert knowledge. Some of these authors
knew Möbius personally and referred directly to his studies. The book Dorfteich als
Lebensgemeinschaft (Village Pond as a Living Community), published in 1885 and
written by Friedrich Junge, is in some sense an offshoot of Möbius’s teaching activi-
ties in teacher training colleges; at the same time it was a book that was to play an
important role in the dissemination and popularisation of a way of looking at
research objects and one that certainly helped to prepare the field for the appearance
of a new discipline called limnology. Thus Dorfteich is not merely a textbook or a
schoolbook,29 a text that might be regarded as a means of establishing certain disciplinary
26 This first volume was followed in 1872 by a second, based likewise on studies – mainly observations
– conducted by Möbius from 1860 onwards in Kiel Bay with his friend, patron and co-editor of
the two volumes, Adolf Meyer-Forsteck. In 1869, at the age of 43, Möbius was appointed
Professor of Zoology.
27 An authorised English version of this work (translated by H.J. Rice) was published in 1883.
28 “Jede Austernbank ist gewissermaßen eine Gemeinde lebender Wesen, eine Auswahl von Arten
und eine Summe von Individuen, welche gerade an dieser Stelle alle Bedingungen für ihre
Entstehung und Erhaltung finden, also den passenden Boden, hinreichende Nahrung, gehörigen
Salzgehalt und erträgliche und entwicklungsgünstige Temperaturen.” Möbius 2006 (1877), p. 75;
see also Reise 1980, Jax 2002, pp. 32 ff.
29 Junge himself notes explicitly that “Village Pond is definitely not intended as a book from which
one can teach” (der Dorfteich absolut nicht ein Buch sein soll, aus dem man unterrichten kann,
Junge 1885, p. VIII).
244
A. Schwarz and K. Jax
facts, theories, and methods by closing the field, creating a sense of “normality”,
reifying accredited knowledge and excluding open questions. Instead, with an eye to
popularization the book additionally opens up a new field of scientific culture, inso-
far as Junge calls for a “deeper study of life in nature” (tieferes Studium des Lebens
in der Natur; Junge 1885, p. IX), advocating that the village pond is an ideal object
for illustrating what we can learn about the laws of nature “out there” in its manifold
forms. In this respect Möbius’s concept of the Lebensgemeinschaft appears to Junge
to be especially salient because it refers to a delimited space which can be examined
and which at the same time facilitates greater knowledge about the whole Earth as a
biocoenosis30: “Now every last corner could be regarded as a world in itself, so that
it later became possible, using such mirror images of the whole, to cast our gaze
upon the Earth as the largest living community”31 (Junge 1885, p. IX).
Other textbooks dealing mainly with ecological issues followed soon afterwards,
but none of them offered this particular mix of educational, scientific, anthropo-
geographical and natural history narratives. Either they were written in a more
popularizing style, or else they were oriented towards an explicitly specialized
readership. The Handbuch der Seenkunde. Allgemeine Limnologie by François-
Alphonse Forel (1841–1912) in 1901 might be regarded as one of the first textbooks
of this latter style in the field of aquatic ecology. The rapidly growing number of
textbooks that appeared over the subsequent three decades points to the disciplinary
consolidation of ecology, particularly of aquatic ecology. This is what Friedrich
Lenz emphasizes in his Einführung in die Biologie der Süsswasserseen (Introduction
to the Biology of Freshwater Lakes) in 1928:
Since hydrology now appears to have reached a certain end point in its initial development,
the time would seem to have come to present its research results and research issues in a
form which does justice to differing expectations, giving both the autodidact and the
research biologist, the teacher and the pupil something to work with. […] This [material]
presents a specific topic out of the overall field of freshwater research and yet is quite
particularly suited to providing an introduction to the research issues in this field. The lake
is virtually paradigmatic for the entire field of hydrobiological research.32
30 The expression “the whole Earth as a biocoenosis” is all the more interesting because the concept
of “biosphere” was introduced at about the same time. Geographer Eduard Suess used it first in
his popular textbook Das Antlitz der Erde (The Face of the Earth) 1883–1909. Although Suess did
not come up with a research programme (this was developed by Vernadsky only from 1926
onwards), the biosphere eventually succeeded in becoming the concept that described the whole
Earth as an organism.
31 “Nun konnte jeder kleine Winkel als eine Welt für sich betrachtet, und später von solchen Spiegelbildern
des Ganzen aus ein Blick auf die Erde als größte Lebensgemeinschaft geworfen werden”.
32 “Da nunmehr die Hydrobiologie einen gewissen Abschluß ihrer ersten Entwicklung erreicht zu
haben scheint, dürfte der Zeitpunkt gekommen sein, ihre Forschungsergebnisse und
Problemstellungen in einer Form zur Darstellung zu bringen, die den verschiedenen Ansprüchen
gerecht wird, die also sowohl dem Autodidakten wie dem forschenden Biologen, dem Lehrer wie
dem Schüler etwas gibt. […] Dieser [der Stoff] stellt zwar ein spezielles Thema aus dem
Gesamtgebiet der Süßwasserforschung dar, ist aber in ganz besonderem Maße geeignet zur
Einführung in die Problemstellung dieses Gebietes. Der See ist geradezu das Paradigma für die
ganze hydrobiologische Forschung.” (Lenz 1928, p. III).
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19 Early Ecology in the German-Speaking World Through WWII
Although Lenz is still addressing a broader audience – the autodidact, the biologist
doing research, the teacher and the student – his textbook is written in a completely
different style from that of Junge’s text. Historical reflections are restricted to a
rather short introduction and concentrate mainly on mentioning the most important
scholars in the field. “These kinds of references give students and experts the feeling
that they are participants in a solid historical tradition. And yet the tradition conjured
up by the textbook in which the scientists think they are participating, actually
never existed” (Kuhn 1988, p. 149). This is how philosopher Thomas S. Kuhn
assessed the role of textbooks, and the “Introduction to the Biology of Freshwater
Lakes” seems to support this statement.
In addition to Lenz’s Biologie der Süsswasserseen there was August Thienemann’s
Limnologie. Eine Einführung in die biologischen Probleme der Süßwasserforschung
(Limnology: An Introduction to the Biological Problems of Freshwater Research)
(1926), the first nominally entitled Einführung in die Limnologie (Introduction to
Limnology) (1930) by Vincenz Brehm, but also, as early as 1909, Das Leben des
Süßwassers (Freshwater Life) by Ernst Hentschel, followed in 1923 by the
Grundzüge der Hydrobiologie (Principles of Hydrobiology), covering both marine
and lacustrian research objects. A book with a more explicitly zoological focus was
Biologie der Wasserinsekten. Ein Lehr- und Nachschlagewerk über die wichtigsten
Ergebnisse der Hydro-Entomologie (Biology of water insects: A teaching and
reference work for the most important results of hydro-entomology) (1934) by H.H.
Karny. In the 1930s, elements of “space and nation” were increasingly present in
the ecological literature too, good examples being Ökologie als Wissenschaft von
der Natur oder Biologische Raumforschung (Ecology as a Science of Nature or
Biological Research of Space) by Karl Friederichs, and also, certainly much less
influential, Der Süßwassersee (The Freshwater Lake) by Fritz Steinecke in 1940.
Thus, in the mid-1920s the first step was taken towards the disciplinary consoli-
dation of aquatic ecology, manifested in the successful establishment of journals,
research laboratories, a body of researchers, a scientific society, and recognition by
the scientific and political establishment.
At about the same time, plant ecology was also beginning to take shape more
clearly. There were above all two books, which became what might be considered
foundational works: The Handbuch der Pflanzengeographie (Handbook of plant
geography, 1890) by Oscar Drude and Pflanzengeographie auf physiologischer
Grundlage (Plant geography upon an ecological basis, 1898) by Andreas
Schimper.33 In the 1920s two other textbook followed which had decisive influence
on plant ecology, namely Pflanzensoziologie. Grundzüge der Vegetationskunde
(Plant sociology: Foundations of phytosociology, 1928) by Josias Braun-Blanquet,
and Einführung in die allgemeine Pflanzengeographie Deutschlands (Introduction
to the general plant geography of Germany, 1927) by Heinrich Walter. Also in animal
33 Both books were widely received also beyond the German language area. According to Tobey
(1981), for example, Drude’s book was a major influence for the development of Frederic
Clements’ ideas of ecology.
246
A. Schwarz and K. Jax
ecology, the 1920s saw the publication of important textbooks, including especially
Friedrich Dahl’s Grundlagen einer ökologischen Tiergeographie (Principles of
ecological animal geography) (1921) and Richard Hesse’s seminal work
Tiergeographie auf ökologischer Grundlage (Animal geography upon an ecological
basis, 1924).
Ecological ideas in applied sciences such as agriculture, forestry and pest
control were promoted especially by Karl Friederichs in two volumes entitled
Die Grundfragen und Gesetzmäßigkeiten der land- und forstwirtschaftlichen
Zoologie (The basic questions and regularities of agricultural and forest zoology,
1930). Despite its supplementary title – “especially of entomology” – these
books were appreciated as a generic contribution to the dissemination and stan-
dardization of ecological methods and concepts. Billed as the “ecological part”,
the first volume deals mainly with theoretical and conceptual questions and
offers a discussion of concepts and laws in ecology, of population biology and
autecological factors, of the interrelations between organisms and their environ-
ment, with special consideration of soil life (edaphon). The second volume, the
“economic part”, introduces the major issues in forestry and agriculture such as
pest control, epidemiology, domesticated animals, breeding and also commercial
regulation. Thus the two books offer a synthesis of an ecotechnologically ori-
ented field34 and of scientific ecology. Friederichs writes: “Recently the entirety
of the applied biological disciplines has been brought together by some in a
‘technical biology’. This designation is intended to stress that we should be
more open to the spirit of invention, technical thinking and technical methods
than we have been to date”35 (1930, Vol. 1, p. 15).
Research Programmes and Networks
This survey of the topology of early German-speaking ecology substantiates the
hypothesis that a distinction was made according to the field of objects on land (or
“in the air”) and in the water, that is, between terrestrial and aquatic ecology. It was
a distinction that simultaneously accepted physiographic conditions as a central
issue for ecology. Furthermore, it confirms that the field of aquatic ecology was the
better organized and institutionalized ecological community, a feature that was also
34 In a general sense, ecotechnology mainly develops local theories and practices and is referred to
as applied research, or “use-inspired basic research”. Good examples of recent ecotechnologies are
restoration ecology, landscape ecology, and industrial ecology. By contrast, ecoscience is character-
ized by the development of more general concepts and theories, such as the competitive exclusion
principle, models of predator–prey relationships, as well as models in ecosystem theory.
35 “Neuerdings wird von einigen die Gesamtheit der anwendenden biologischen Disziplinen als
‘Technische Biologie’ zusammengefasst. Durch diese Bezeichnung soll betont werden, daß
mehr als bisher Erfindergeist, technisches Denken und technische Methoden bei uns Eingang
finden sollten”.
247
19 Early Ecology in the German-Speaking World Through WWII
recognized at the time.36 However, the early establishment of common ground in
aquatic ecology is reflected not only in the founding of journals and the compiling
and disseminating of reference works and textbooks. Terrestrial ecologists also
joined together institutionally and epistemically much later. While botanists were
able to close the gap relatively quickly, zoologists formed much later into fields
such as animal or population ecology – in contrast to the situation, for instance,
in the US or the UK. One might argue that research and teaching in the classic
disciplines of zoology and botany were oriented towards other scientific ideals and
programmes, which were too predominant to allow for autonomous ecological
disciplinary activities. By contrast, in the relatively new and institutionally still
unstructured field of limnetic systems, zoologists and botanists could situate their
inquiries within the subject matter of ecology much more easily.
The different traditions of early German-speaking ecology highlight perfectly
the main roots from which ecology originated. We find an important physiologi-
cal or autecological research tradition, which around 1900 merged partially with
what we might call the classificatory or natural history tradition to become what
was then termed the “ecological biogeography of plants and animals”. At the
same time, we find a holistic or systemic tradition which, in different manifesta-
tions, extended beyond the biological aspects of nature to encompass a descrip-
tion of whole entities constituted both by biotic and abiotic elements of nature
(Jax 1998).
These traditions were also taken up, albeit with differing emphases, in an epis-
temological approach that offers to characterize ecology according to three concep-
tual templates, or basic conceptions, namely “microcosm”, “niche” and “energy”.
These allow for a systematic description of the often disparate theories and nar-
ratives in ecology, such as those concerning the concepts of population, accom-
modation or fitness, ideas about the transport of nutrients and organic matter, but
also models designed to simulate, for instance, the productivity of a system, or
predation between certain units. The notion of “microcosm” is based roughly on
a romantic philosophy of nature and approximates to the physiognomic tradition,
while “energy” comes close to the holistic tradition and is basically the most
physical one: like energy in physics, organic matter in ecology becomes the basic
dimension of properties and agency and enables the integration, for instance, of
the living and non-living in a single system – an ecosystem. Finally, the basic
conception of “niche” concentrates on concepts that were developed within the
36 Thienemann notes retrospectively (1939, p. 13): “As the first of three sub-sciences, limnology
has returned to its universal ecological objective and pursues it deliberately (after plant ecology
had developed at least partly in the same direction)” (“Die Limnologie hat sich als erste der drei
Teilwissenschaften – nachdem sich die Pflanzenökologie zum Teil wenigstens in der gleichen
Richtung entwickelt hatte – auf ihre allgemeine ökologische Zielsetzung besonnen und verfolgt
sie bewußt.”) However, Thienemann certainly did not deny that plant sociology had a tradition
stretching back longer than limnology. But in order to genuinely be doing ecology – to be “bio-
coenotics” – the terrestrial zoologists and botanists would have to join together more closely, as
was the case, he claimed, in aquatic ecology (1925, p. 75 f.).
248
A. Schwarz and K. Jax
framework of evolutionary biology to characterize the relationship between
individual organisms, species and their environments; as such, it is similar to the
physiological tradition.37 These three basic conceptions became decisive for the
establishment and stabilization of ecology as a scientific discipline. According to
Schwarz, ecology developed as a pluralistic endeavour from the beginning, while
each of these basic conceptions ultimately guides more implicitly the building of
hypotheses and concepts (see Chap. 8). Following philosopher Imre Lakatos, they
are hidden in the “hard core” of a research programme, where these unverbalised
ideas of nature irreducibly influence theory building. Another philosopher,
Gernot Böhme, proposes from a phenomenological perspective that we talk of the
character of nature in analogy to the way we talk about the character of people.
Both concepts may be helpful in grasping the idea of the basic conceptions that
are to characterize a particular field of ecological knowledge embracing prac-
tices, narratives and theories about living beings in both aquatic and terrestrial
environments.38 In the following, both conceptualizations are used complemen-
tarily to one another in order to chart the field of early ecology in the German-
speaking world.
Ecological Water Affairs
From the eighteenth century onwards, geomorphological and hydrological descriptions
of rivers, limited regions of the sea, and lakes emerged.39 However, these bodies of
water were merely perceived as “water deserts” – devoid of life – and it was not
until about 1870 that they were widely acknowledged as an environment that
played host to a plethora of macro- and microorganisms. A number of concepts
were offered to describe these lakes, suddenly full of life and attractive objects to
be investigated by zoologists, botanists, and microbiologists. One of the first concepts
to circulate in the biological community was that of “the lake as a microcosm”;
virtually simultaneously the lake was introduced as an “organism”, an “island” and
37 The relation between the pattern of the three “traditions” and the three “basic conceptions” is
certainly not always equally close but rather involves a roughly drawn analogy and would require
further investigation. However, the two analytical patterns make even more clear the divide
between terrestrial and aquatic ecology in early German-speaking ecology. To mention just one
important difference, the use of the word “organism” varies between the different traditions. While
the “physiological tradition” is closely related to autecology and thus focuses on the single organ-
ism, the basic concept “energy” includes models and metaphorical conceptualizations of the
organism which refer to the physiology of a lake or parts of the ocean.
38 See Schwarz Chap. 8 this volume. An example focusing on aquatic microbiology is given in
Psenner et al. 2008.
39A neat hydrological description of Lac Leman was provided as far back as 1779 by the famous
Swiss naturalist Horace-Bénedict de Saussure (1740–1799), and of Lake Constance by David
Hümlin in 1783. Rivers were also subject to geomorphological and hydrological measurements at
the time (Schwarz 2003a, pp. 116 ff.).
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19 Early Ecology in the German-Speaking World Through WWII
a “geographic individual”.40 The Swiss naturalist François-Alphonse Forel wrote of
the lake as a system, a conceptualization that was to become crucial not only in
turning limnology into a scientific discipline (1886) but also in providing a first, if
still somewhat hazy, blueprint for a systemic understanding of the lake. Forel
depicted the lake as a laboratory model for large-scale phenomena also happening
“in the immensity of the ocean” yet not accessible to scientific analysis on this
scale, concluding: “It is much easier to study a lake than an ocean” (Forel 1896).
Forel paved the way for consideration of the lake as an experimental system, looking
at the lake as a system in which organisms interact, certain functions are processed,
and an exchange and circular flow of materials occurs. The lake is constituted as a
system in which abiotic and biotic elements are drawn closely together. In this, the
“organic substance” is subject to constant “transformations and migrations”, is
“incarnated” in living organisms and released again. “The organic substance returns
time and again to the great provision house, the lake, be it in the form of animal
secretions such as carbon dioxide, urea or other products of the animal metabolic
processes, or after the organisms have died as products of decay. […] The organic
substance released provides an inexhaustible, continually renewed supply from
which animals and plants renew their structure […]. This microcosm, to use the
term introduced by Prof. S.A. Forbes, would be sufficient unto itself in the long
term, even if it were to become completely isolated from all surrounding media”41
(Forel 1901, pp. 237, 239).
At the end of his handbook Forel proposes a “programme for limnological inves-
tigations” in which the biological study of the fauna and flora of a lake is just one
of nine headings, the last heading in a long list of issues dominated mainly by
physico-chemical and geoscientific perspectives. The programme represents an
approach, which focuses mainly on research concerning the cycles of elements in
lakes, the functionality of organisms and organic substance in these dynamic cyclical
processes, as well as the flow of energy through aquatic systems. Following the
conceptual scheme mentioned above, these elements are representative of the template
40 Forel 1901; Otto Zacharias, Skizze eines Spezial-Programms für Fischereiwissenschaftliche
Forschungen (Sketch of a special programme for scientific fisheries research) in: Fischerei-
Zeitung 7 (1904), pp. 112–115; Über die systematische Durchforschung der Binnengewässer und
ihre Beziehung zu den Aufgaben der allgemeinen Wissenschaft vom Leben (On the systematic
exploration of the inland waterways and their relationship to the tasks of the general study of life)
in: Forschungsberichte der biololgischen Station Plön 12 (1905), pp. 1–39; Das Süßwasserplankton
(Freshwater plankton). Leipzig: Teubner 1907; the classic paper The lake as a microcosm by
American entomologist Forbes was published in 1887.
41 “Die organische Substanz kehrt immer wieder in die große Vorratskammer, den See, zurück,
sei es in Form von tierischen Sekretionen, wie Kohlensäure, Harnstoff und anderen Produkten
der tierischen Verbrennungsvorgänge, oder nach dem Tode der Organismen als Produkte der
Verwesung. […] Die gelöste organische Substanz stellt einen unerschöpflichen, stets erneuerten
Vorrat dar, aus dem Tiere und Pflanzen das Material zu erneuertem Aufbau entnehmen […]
Dieser Mikrokosmos, um den von Prof. S.A. Forbes eingeführten Ausdruck zu gebrauchen,
würde sich selbst auf lange Zeit genügen können, auch wenn er gegen die umgebenden Medien
völlig isoliert würde”.
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“energy”, expressed in concepts relating to the productivity of lakes (Waldemar
Ohle and many others), to theories describing lakes as circular causal systems, and to
systems ecology in general inspired by cybernetics, in which the analysis of bioener-
getic transfers became a central research question (see also Part 10); the latter pro-
gramme was developed largely in the US, at least to begin with (mainly by George
E. Hutchinson, Raymond Lindeman, Howard Odum & Eugene Odum). By contrast,
the emphasis in the German-speaking ecological world was on the productivity
approach which, in a certain sense, referred to the theoretical idea of an “outer physi-
ology”, looking at the physiology of the “organism lake” (or ocean), while at the same
time being geared in a less theoretical way towards studies in farming and fisheries.
Forel’s “limnological programme” had been spelled out for the first time in
1886 in a French publication and was taken up soon after that by a number of natu-
ralists. These activities also had institutional backing, namely from the limno-
logical commission of the Swiss naturalist society. What started out as a mere
observation programme about the freezing of Swiss lakes which brought together
laymen and scientists eventually culminated in systematic long-term observations
at Lake Zurich and nearby Lake Walen, at Lake Constance and, in particular, at Lake
Lucerne. The latter study was unique at the time, not only because of the length
of time over which data were collected, but also because of the density of the inter-
vals: this ecological long-term study – probably the first of its kind – began in
1896 and lasted over a period of roughly 5 years.42 The number of samplings per
year was stipulated at the start of the campaign, and a precise description of sites
as well as detailed methods were provided. Despite the different disciplinary and
institutional background of the naturalists who participated, an attempt was made
to systematize and standardize sampling, methods and instruments. The results
were published in a series of papers, starting in 1900 and ending in 1917.43
Again, for the first time in the short history of limnetic ecology, chemical, physical
and biological data were related to one another, facilitating an integrative view of the
interrelationships, say, between temperature, quantity of phytoplankton, organic
substance and carbon stored in calcium carbonate. Ultimately, it was not only the
42 The programme was described in detail by Xaver Arnet, a secondary school teacher in Lucerne,
and was published in the communications of the natural history society of Lucerne in 1895. It also
included a call to all members of the society to participate in the data collection, which again documents
the institutional openness of the field (“Programm zur limnologischen Untersuchung des
Vierwaldstätter Sees. Programm für den physikalisch-chemischen Teil”. In: Mitteilungen der
Naturforschenden Gesellschaft Luzern 1, pp. 1–16).
43 To list just a few of these publications: Bachmann, Hans (1904). Das Phytoplankton des
Süsswassers (Freshwater phytoplankton), in: Botanische Zeitung 62, 82–103; Burckhardt,
Gottlieb (1900). Quantitative Studien über das Zooplankton des Vierwaldstättersees (Quantitative
Studies on the zooplankton in the Vierwaldstätter lake), in: Mitteilungen der Naturforschenden
Gesellschaft Luzern 3, pp. 129–411, 686–707, 414–434; Amberg, B. (1904). Limnologische
Untersuchungen des Vierwaldstättersees (Limnological studies of the Vierwaldstätter lake), in:
Mitteilungen der naturforschenden Gesellschaft Luzern 4, pp. 1–142; Nufer, W. (1905). Die
Fische des Vierwaldstättersees und ihre Parasiten (The fish of the Vierwaldstätter lake and their
parasites), in: Mitt. naturf. Ges. Luzern 5: 1–232.
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research practice – the modality of acting in and on the lake – but also the style of
representation that changed profoundly during the course of the project. Forel had
already invented the “depth ordinate” (Tiefenordinate) by moving the abscissa to the
top and depicting measured data downwards along the ordinate, thereby representing
measuring in the lake into depth.44 This type of diagram was also taken up in the
Lake Lucerne studies and used in various representational modes while experimenting
with time scales, multiple sampling sites and parameters (Fig. 19.2).
Fig. 19.2 Burckhardt uses the depth ordinate to depict the distribution of zooplankton over time.
What makes the figure rather puzzling for contemporary, visual practice is the fact that he includes
several sites in the same coordinate space without marking them (1900, p. 424)
44A more detailed discussion on the impact of a visual language in aquatic limnology is given in
Schwarz (2003b).
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All these activities were considered to be rather avant-garde in limnological
research, causing German plankton specialist Otto Zacharias to note: “Switzerland can
lay claim to the fame of being the classical country not only of lakes but also of lake
exploration”45 (1888, p. 214), a statement he still stood by roughly 10 years later.
Zacharias was not only an admirer of Swiss aquatic research, and especially of
Alphonse-François Forel, but was himself quite a well-known plankton specialist.
He was interested in collecting plankton species, describing their manifold forms
and their collectivization (Vergesellschaftung). More specifically, he took up
Forel’s main area of interest, namely to discover more about dead and living
organic substance. However, for Zacharias this was just one among many other
approaches. He was convinced that we need to discover, first of all, the many species
not yet known to us, and also to learn much more about the species we do already
know. Accordingly, a significant proportion of his publications is devoted to the
description of plankton organisms and their behaviour in different lakes, seasons,
and plankton communities, although he never delved too deeply into aspects of
quantification.46 In the thoroughly vitalized world of Zacharias, organisms were
bound up within superordinate webs of connection. Zacharias’s nature rests in a
stable balance which prescribes certain functions to its organisms. There is no contra-
diction between this view and one that accepts the existence of a law of metabolism
for the organic substance, a law which, he says, can be discovered in animal and
plant communities, as the famous August Weismann had “astutely” established in
his treatise on animal life in Lake Constance (1905, p. 31). However, in his description
of the way this metabolism worked – which elements in it supposedly function in
what way – Zacharias gets only a little further than the description given by
Weismann about 30 years previously.47
45“Die Schweiz […] darf den Ruhm für sich in Anspruch nehmen, das classische Land nicht bloß
der Seen, sondern auch der Seendurchforschung zu sein”.
46Zacharias was surely one of those nature researchers whom Lauterborn somewhat derogatorily
referred to as “high climbers” – quoted thus by American limnologist Henry B. Ward in Science:
“And what Lauterborn said five years ago is even truer today in the light of our more extended
experience: ‘For the question as to the distribution of organisms, the methods so cherished even
up to the present day of fishing in the greatest possible number of lakes (which recalls, in many
respects, the chase after summits on the part of our modern high climbers – Hochtouristen!), really
have only limited claim to scientific value, since through them but a very incomplete picture of
the faunal character of a water basin can be obtained.” (1899, p. 499).
47Albeit this was a study which Weismann himself had presented neither as a research report nor to
a scientific audience but rather as a “readily accessible talk” which he had “held in the main hall of
the University of Freiburg in front of an audience consisting largely of ladies” (Weismann 1877, p. 3).
The fact that nonetheless reference was repeatedly made to this talk given by an established scientific
authority serves to illustrate the extent to which the status of aquatic ecology was still uncertain
around the turn of the century. Another point of reference to do with the content of the talk may
well have been that Weismann assumed the transmutation of species through the influence of external
conditions and thus supported a Lamarckian rather than a Darwinian conception of evolution. Not
until his later “genetic” studies did Weismann come to adopt a Darwinian position.
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19 Early Ecology in the German-Speaking World Through WWII
The descriptions given by Zacharias which proved to be relevant were, at best,
those with which an otherwise indeterminate “whole” is replaced by the lake’s
“household of nature”. This Naturökonomie (natural economy) can most certainly
be measured and its productivity established in quantitative terms – an endeavour
in which Zacharias was building on a successful research programme.
However, like many of his contemporaries in early ecology, Zacharias basically
specialized in a scientific style geared towards natural history, that is, dedicated to col-
lecting, organizing, putting in order and eventually establishing a physiognomic
method. This method was common in geobotany, plant sociology and plant geography,
including the description of aquatic plants; but it was also used to develop a typology
of lakes and landscapes based mainly on either gestalts or typical animals and plants.
To return to the conceptual pattern proposed earlier, all this was typical of the basic
concept of “microcosm”. The following quote may serve to demonstrate that Zacharias,
and many other scientists as well, were devoted to a romantic vision of nature; in it, he
vigorously opposes those positions in which nature is claimed to be a machine or some-
thing gruesome and threatening: “There is also a biological optimism in which nature,
when viewed through its eyes, is most definitely not some eternally devouring, eternally
ruminating monster, but is rather a goddess that conjures forth ever new inexhaustible
life from death and decay, and whose reign challenges our admiration all the more as
we become more intimately familiar with it through serious study”48 (1907, p. 9).
Literary Genre and Data Representation
Up until the end of the 1920s, taxonomy and data collection were still a dominant
part of most research activities in the aquatic realm. In a way, the review given by
Kurt Lampert in Das Leben der Binnengewässer (The Life of the Inland
Waterways),49 a handbook in three editions expanded continually between 1899 and
1925, was a preliminary culmination of this first period of collection in aquatic
ecology. Another project, even more encyclopaedic, was a handbook on plant ecology,
Die Lebensgeschichte der Blütenpflanzen in Mitteleuropa – spezielle Ökologie der
Blütenpflanzen Deutschlands, Österreichs und der Schweiz (The life history of flowering
plants in Central Europe – special ecology of the flowering plants of Germany,
48 “Es gibt auch einen biologischen Optimismus, mit dessen Augen angesehen die Natur durchaus
kein ewig verschlingendes, ewig wiederkäuendes Ungeheuer ist, sondern vielmehr eine aus Tod
und Verwesung immer neues unerschöpfliches Leben hervorzaubernde Göttin, deren Walten
unsere Bewunderung um so mehr herausfordert, je genauer wir uns mit ihm durch ernste Studien
bekannt machen”.
49 Das Leben der Binnengewässer was first published in 1899 by Kurt Lampert (Leipzig: Tauchnitz),
the second edition in 1910 (by then it had grown from 591 to 856 pages), while the third edition of
1925 (892 pages) was edited by R. Lauterborn, V. Brehm and A. Willer. The recent project Die
Süßwasserfauna von Mitteleuropa (Freshwater fauna of Central Europe), established by A. Brauer
and encompassed in 21 volumes edited by J. Schwoerbel and P. Zwick between 1985 and 2000, also
illustrates the fundamental importance of such encyclopaedic projects for ecological research.
254
A. Schwarz and K. Jax
Austria and Switzerland), edited by Carl Schröter, Oskar von Kirchner and E. Loew.
The first volume was published in 1908 and the last in 1942. In animal ecology,
again, classification from an ecological vantage point occurred rather late and was
not initially manifested in the form of a handbook or series50; instead, this kind of
knowledge appeared to be organized mainly in journals such as the Zoologische
Jahrbücher. Abteilung für Systematik, Ökologie und Geographie der Tiere
(Zoological Yearbooks. Department for Systematics, Ecology and Geography of
Animals) (1926–1994), the Zeitschrift für Ökologie und Morphologie der Tiere
(Journal of Ecology and Morphology of Animals) (1924–1967); around this time
also, Richard Hesse published his monograph Tiergeographie auf ökologischer
Grundlage (Animal Geography upon an Ecological Basis) (1924).
A similar type of classificatory work was the Handbuch der biologischen
Arbeitsmethoden (Handbook of biological study methods). This – as its subtitle states –
was a “comprehensive compendium of methods which embraces the entire scientific
field of study and research”. The handbook, edited by Emil Abderhalden, was begun in
1920 and covered a whole wide range of disciplines, from geology and palaeobiology
to physiology and medicine; it also offered a detailed account of physical and chemical
methods, instruments and materials. Einar Naumann, August Thienemann, Friedrich
Lenz and others contributed to “the methods of freshwater biology” as part of the section
entitled Methoden der Erforschung der Leistungen des tierischen Organismus (Methods
for Researching the Functions of the Animal Organism, 1926).
Another basic literary genre were monographs on lakes and rivers. In some sense,
these monographs fit very well with the ecological perspective regarding the indi-
viduality of its objects; the idea was to look at the lake as a geographical individual
or as an organism that is born and then dies off. Both concepts were fairly common
and were consolidated conceptually in the system of lake types, which encouraged
aquatic ecologists to regard lakes – including their living and non-living parts – as
evolving units. The three-volume Le Léman: Monographie limnologique by François-
Alphonse Forel (1892–1904) was probably the most influential work here, function-
ing as a kind of blueprint. Of course, it was written in French, but it was well received
nevertheless by the German-speaking community. This was almost certainly because
Forel also published in German and, furthermore, was in close correspondence with
his German-speaking colleagues. Other limnological monographs of a lake included
the Würmsee (1901) and the Ammersee (1906) by Willi Ule and, to some extent, the
Vegetation des Bodensees (Vegetation of Lake Constance) (1896, 1902) by Schröter
and Kirchner. Insofar as the latter contains first a general scientific characterization of
Lake Constance, however, the focus on botanical objects – plankton and macrophytes
– is quite explicit. Running water was also the subject of monographic inquiries, most
prominently the detailed description of the river Rhine by Robert Lauterborn, starting
with several publications on “the geographical and biological structure of the river
Rhine” between 1916 and 1918, based on and followed by studies of the Upper
50 Die Ökologie der Tiere (Ecology of Animals) by Fritz Schwerdtfeger, published in three volumes
between 1963 and 1975, might be regarded as a belated first series in animal ecology, comparable
in some sense to Lampert’s Binnengewässer.
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Rhine and Lake Constance. In 1930 a first part of the monograph Der Rhein:
Naturgeschichte eines deutschen Stroms (The Rhine: Natural history of a German
river) was published, followed by a second and third part in 1934 and 1938.51
Classifying Lakes and Rivers – Relating Type and Process
Also part of the activities engaged in around classification was the development of a
system of lake types which, to begin with, followed rather geographical and/or purely
botanical or zoological criteria: Coregonenseen und Zanderseen (whitefish lakes and
pikeperch lakes), Dinobryonseen und Chlorophyceenseen (dinobryon lakes and
chlorophyceae lakes) are just some of the proposed types. Starting with more specific
studies on the physiographic character of a lake, naturalists realized that the distribution
and composition of organisms depends critically on the chemical and physical param-
eters of lake water. This was the start of a very influential programme, which sought
to combine causal relations with physiognomic traits. The study of lake types absorbed
a number of German and Scandinavian researchers and served as an overall conceptual
framework in which much empirical research was embedded. While Thienemann and
American limnologists Edward A. Birge (1851–1950) and Chancey Juday (1871–1944)
conducted similar studies, they drew very different conclusions from the results. Both
were interested in the relationship between physico-chemical conditions and the abun-
dance of plants and animals in a lake. Birge and Juday concentrated mainly on the
seasonal and diurnal dynamics of an individual lake, taking a physiological perspective
on the lake as a system, whereas Thienemann tried to incorporate his findings into the
overall scheme of a conceptual system of lakes. In his classificatory approach, he
combined geographical zones, animal indicators and physical and chemical features –
such as the thermocline or carbon concentration – and came up with a complex typology
that was ultimately more confusing than illustrative, as the following comment rather
unintentionally reveals: “The Chironomus lake had now acquired the designation
‘Baltic lake’, since it predominated in that area, whereas the Tanytarsus lake was called
a ‘subalpine lake’; neither was intended as a geographical term”.52
51 The story of this monograph is interesting in itself, but would go beyond the scope of this account;
for more detail, see in RegioWasser e.V. (ed) (2009). 50 Jahre Rheinforschung. Lebensgang und
Schaffen eines deutschen Naturforschers Robert Lauterborn (1869–1952). Freiburg: Lavori Verlag.
52 “Der Chironomussee hatte nunmehr die Bezeichnung ‚baltischer See’ erhalten, da er in diesem
Gebiet vorherrschte, während der Tanytarsussee ‚subalpiner See’ genannt wurde; beide sollten
keine geographischen Begriffe sein”. This comment was made by Friedrich Lenz, former assistant
to Thienemann, at the IV. Hydrobiologische Konferenz der Baltischen Staaten in Leningrad,
September 1933. The title of his talk was “Das Seetypenproblem und seine Bedeutung für die
Limnologie” (The problem of lake types and its significance for limnology). At the time, the
terminology of lake types had already shifted to the terminology of poly-/eu-, meso-, and oligo-
trophy. Even Thienemann himself had given up the geographical/zoological terminology. As early
as 1921 he subsumes the Baltic type within the eutrophic type: “Vor allem im Flachland des
Baltikums verbreitet, aber auch in den Alpen vertreten. Häufig in Nordamerika” (Common above
all in the Baltic plains, but also found in the Alps. Frequently in North America); 1921, p. 345).
256
A. Schwarz and K. Jax
At about the same time, Swedish limnologist Einar Naumann (1891–1934)
developed his “regionale Limnologie”, the aim of which was likewise to conduct a
“causal study of the distribution of types of water bodies in general with their specific
world of organisms on Earth” (kausale Studium der Verbreitung der Gewässertypen
im allgemeinen mit ihrer speziellen Organismenwelt auf der Erde; 1923, p. 75).
From the very beginning Naumann concentrated on describing the productivity of
lakes and, as early as 1918, he proposed an interesting typology that remains little
known and indeed is not as interesting for the concepts proposed in it as for the
method used. First and most notable, it was not based solely on empirical studies
from different types of ponds at the fisheries experiment station at Aneboda; rather,
Naumann drew his results from an experimental situation outside the laboratory.
This approach was rather unusual at the time, as the following comment by
Thienemann may illustrate: “The method to be applied in tackling our problem
[establishing types of lake] is that of comparative observation in nature; experimen-
tal study of individual factors can – as indicated above all by Einar Naumann’s
studies – provide clarification and greater detail”53 (1921, p. 344).
In his experimental system Naumann distinguished between a Naturtypus (natural
type) and a Kulturtypus (cultural type), indicating mainly the depth of intervention:
the natural type refers to ponds which “are neither given fertilizer nor used as feeding
ponds”, while the cultural type are ponds “which are either given fertilizer or serve
as feeding ponds.”54 He proposes four types and comes to the conclusion that it is
possible to consider “the production of phytoplankton, determined both quantita-
tively and qualitatively, as by far and away a highly accurate indicator of milieu”
(die Produktion an Phytoplankton, quantitativ und qualitativ ermittelt, mit grossem
Vorteil als einen sehr scharfen Milieu-Indikator verwerten [sic!], 1918b, p. II). He
urgently cautions his readers, however, against using such an “algological crediting
method” for purposes of fisheries biology as well – the link between the production
of fish meat and that of phytoplankton, he says, is too complicated. In the following
year he comes up with the concepts of eutrophic and oligtrophic lakes55 and, two years
later, comments: “When Kolkwitz and Marsson first analyzed systematically and in
a modern way the effect of organic fertilizers on water (1908, 1909), the system of
saprobes was established on this basis. Depending on the degree of pollution of
the water, these latter were allotted to the zones of polysaprobes, mesosaprobes and
oligosaprobes. […] Now we might well wish to ask to what extent the system
53 “Die Methode, die bei der Bearbeitung unseres Problems [der Aufstellung von Seetypen] anzuwenden
ist, ist die vergleichende Beobachtung in der Natur; experimentelles Studium einzelner Faktoren
kann, wie vor allem Einar Naumanns Untersuchungen zeigen, Klärung und Vertiefung bringen”.
54 Naumann 1918b, p. II. The quotes originate from a paper published in Swedish; only the sum-
mary is in German, and this again is contained “only in the publisher’s offprints”. Naumann used
the biological laboratory in Aneboda for a number of field experiments and for what we today call
mesocosm experiments, most likely beginning in 1916.
55 Naumann points out that “eutroph” and “oligotroph” had been already used in 1907 by C.A.
Weber in a study on swamps in Northern Germany (Steleanu 1989, p. 391).
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19 Early Ecology in the German-Speaking World Through WWII
proposed by myself is really very helpful. In order to settle this question, it is
necessary to analyse the reciprocal relationships between these ecological systems –
which, of course, may be used quite independently of each other. […] (1) The
physiological system proposed by me serves the purpose of a pure analysis of the
factors determining production each for itself. (2) In contrast to this, the system of
saprobes works with the standard analysis of water”56 (1921, p. 19 f., emphasis in
original). In conclusion, Naumann proposes a “pure analysis” of the “special factors”
in order to investigate ecological systems scientifically using the “trophy stan-
dards”, while the “standard analysis” indicating the saprobial index is appropriate
for applied purposes. He expects that the “milieuspectra” of the different special
factors, for instance the N and P household,57 should then be useful for a “regional
mapping of the various sub-spectra” and thus for an evaluation of the “overall biology
of waterbodies” (1921, p. 20).
“Physiology” in Aquatic Ecology
The trophic system was quickly adopted in the 1920s, and the same applies for the
“milieu-spectra”: “Naumann’s notion of ‘milieu-spectra’ has proven to be thoroughly
stimulating and fruitful in limnology”58 commented Thienemann in his review of
the “nutrient cycle in water” (1927, p. 43).
Naumann used his method mainly to present the “lake” – however technically
modified – as an ecological system which can and must be described in terms of
physiological functions and with respect to its physico-chemical conditions; however,
this same method was also used to represent the milieu standard (or milieu needs) of
a single species. “Physiology” in aquatic ecology could thus refer to the physiology
of a whole lake or to the physiology of a single organism (Fig. 19.3).
56 “Als Kolkwitz und Marsson zuerst (1908, 1909) die Einwirkung von organischen Dungstoffen
auf das Wasser systematisch in moderner Weise analysierten, wurde auf diesem Grund das System
der Saprobien begründet. Je nach dem Verschmutzungsgrad des Wassers wurden dieselben in den
Zonen der Poly-, Meso- und Oligosaprobien eingereiht. […] Die Frage dürfte indessen gestellt
werden können, inwieweit das von mir vorgeschlagene System wirklich weiter führt. Zur
Erledigung dieser Frage ist eine Analyse der gegenseitigen Verhältnisse dieser ökologischen
Systeme – die selbstverständlich ganz unabhängig voneinander gebraucht werden können –
erforderlich. […] 1. Das von mir vorgeschlagene physiologische System bezweckt eine
Reinanalyse der produktionsbestimmenden Faktoren jeder für sich. 2. Das System der Saprobien
arbeitet im Gegensatz hierzu mit dem Durchschnittstandard des Wassers”.
57 The special factors are gases, temperature, or mineral nutrients, such as calcium carbonate,
phosphoric acid, nitrate, but also humin. Temperature, each gas or mineral nutrient has its own
budget with a specific spectrum which is also classified in three realms: polytrophy, mesotrophy
and oligotrophy (1921, p. 5 f.). Each lake can be evaluated on the basis of these spectra of the
budget of each important chemical (or physical) factor.
58 “Naumanns Gedanke der ‚Milieuspektren’ hat sich in der Limnologie als überaus anregend und
befruchtend erwiesen”.
258
A. Schwarz and K. Jax
This was also taken up in other parts of ecology. Zoologist Richard Hesse, in his
book on animal ecology, talks about “the optimal conditions of a biotope” when the
best possible number of species appears (1924, p. 18). The concept of “milieu standard”
and, similarly, “partial ecological spectrum” also come close to today’s concept of
the environmental needs of a species or the distinction between physiological and
ecological optimum.
For a number of limnologists, this kind of research did not go far enough in
terms of physiological and morphological analysis of single organisms and ecological
investigation of individual populations. Richard Woltereck was one of the scholars
who deplored the fact that limnologists were interested only in “general questions of
‘production biology’, of regional limnology, of types of water bodies, ‘milieu-spectra’
etc. […] The reason for this phenomenon lies partly in the low esteem in which
XVIII. XVII.
Poly
_
troph.
Humos meso
_
troph.
Humos Oligo
_
troph.
Humos
Humus
N
+
P
10000
bis
bis
bis
bis
1000
100
10
1
54321
XII. Z. XI. VIV
Thermik
O2
N + P
Humus
PH
CaO
Polytypus Mesotypus Oligotypus
a
b
XXI. X.
Fig. 19.3 (a) Depicted here are the “milieu needs” of Holopedium gibberum, a very common
zooplankter which Thienemann had described in detail in the journal Zoomorphology in 1926
(Thienemann 1927, p. 43). (b) Hans Utermöhl offers a different, more precise representation of a
diatom (Cyclotella comta), speaking of a “partial ecological spectrum” (ökologisches Teilspektrum)
while commenting that the species need not necessarily occur at every line or level. “However,
they may occur there”, writes Utermöhl in his Limnologische Phytoplankton-Studien of 1925
259
19 Early Ecology in the German-Speaking World Through WWII
‘laboratory work’ is held by those hydrobiologists who are unable to think physi-
ologically, and partly also, one can assume, in the awkwardness of such work,
which is nevertheless indispensable if hydrobiology is to achieve its objective as a
science”59 (1928, p. 543).
This statement is explicitly directed against Carl Wesenberg-Lund (1867–1955),
and presumably also against August Thienemann. At the very least, this setting up
of different fronts appears to be a polemical strategy as far as the implied concept
of “milieu-spectra” is concerned, which – as shown above – undeniably reflects a
physiological perspective, was applied to individual organisms and was also based
on practical experimental work.
Woltereck was clearly after an experimental approach in his ecological research:
He set up a laboratory at the Biologische Station in Lunz60 and later a private labora-
tory in Seeon in which cladocerans served as laboratory organisms to investigate
morphological, physiological and autecological questions. In addition, though, he
was interested in field experiments: he exposed Daphnia cucullata in Lake Nemi
(Italy) as an alien species (indigenous in Denmark), and he organized a survey of
the biological repopulation of the same lake after it had been exhausted from water
completely.61 His in situ studies at Lake Lunz resulted in a model, the so-called
Nahrungs-Zehrungs-System (N/Z-system, food-consumption system), to describe a
three-part ecological system consisting of (1) several species of nannoplankton,
(2) a population of Daphnia, and (3) “summer shoals of whiting” (Weißfische)
(1928, p. 544). Before presenting the results of his study, he pointed out that the
whole research design was merely a frame and that the main work had yet to be
done; moreover, the empirical data he presented were indeed rather flimsy. Even so,
he believed that the results of the study showed not only that the “ecological balance
of the population numbers” (ökologische Gleichgewicht der Volkszahlen, 1928,
p. 548) had been maintained, but that this situation of a system equilibrium repre-
sented law-like behaviour and could be described using the model N : Z = K Z : PN,
or, in words: the number of daphnids is in proportion to the number of fish as the
daily consumption of daphnids by a fish is in proportion to the daily reproduction
of daphnids. Woltereck sought to pursue an ecology based on laws or law-like
relationships in order to explain the behaviour of populations, the differences in
lake types, and the harmony of a well-shaped system in general. However, his
emphasis on explanation and laws in ecology was always accompanied by a strong
59 “die allgemeinen Fragen der ‘Produktionsbiologie’, der ‘regionalen Limnologie’, der
‘Gewässertypen’, ‘Milieuspektren’ usw. […] Die Ursache dieser Erscheinung liegt zum Teil in der
Geringschätzung der ‘Laboratoriumsarbeit’ seitens solcher Hydrobiologen, die nicht physiolo-
gisch denken können, teilweise wohl auch in der Unbequemlichkeit solcher Arbeiten, deren
Inangriffnahme gleichwohl unentbehrlich ist, wenn die Hydrobiologie ihr Ziel als Wissenschaft
erreichen will”.
60 Despite being the station’s first director, Woltereck never actually did any administrative work
there, because he took up a position in Leipzig and later also in Ankara. Franz Ruttner was initially
his deputy director, taking on the role of director of the station proper in 1908 (until 1957).
61 Remarkably enough, this draining was done for the purpose of an archaeological project of national
importance that was to recover the galleys said to have been built in ancient times by Caligula.
260
A. Schwarz and K. Jax
commitment to the existence of an “autonomous law” and an “inner indiosyncratic
activity” of the organism – a view in which he essentially finds himself in agreement
with Ludwig von Bertalanffy’s biological holism (1940, p. 476). This belief in the
autonomy of each individual might also be seen as substantiating his opposition to
a type of explanation in biology requiring a supra-individual unity or superordinate
power – such as Thienemann’s cipher of the lake as “representing the unity of
biotope and biocoenosis” (1925, p. 595).
The Physiological Programme in Terrestrial Ecology
The physiological (or autecological), approach in early ecology was, as far as
terrestrial zoology and botany are concerned, basically a Darwinian, or at least
adaptationist, one. This tradition has remained rather neglected within the history
of ecology,62 although it has been highly influential for the formation of ecology,
both in Europe and in the United States. Taking up evolutionary theory, especially
in the wake of Darwin’s Origin of Species (1859), both botanists and zoologist
strove to view organisms and their morphology in terms of their adaptedness to
their environment. In its early stages this research remained a minor undercurrent
of biological research, the mainstream of which was focused on morphology and
physiology (as strictly separate fields) in the laboratory. The main type of
research spawned by evolutionary theory during the second half of the nineteenth
century was phylogenetic morphology, which focused on an analysis of questions
relating to taxonomy and developmental biology. However, a group of younger
scientists also found their way back from the lab to organisms in their natural
habitat. As was the rule during the early years of terrestrial ecology, this occurred
separately in zoology and botany and also with varying degrees of success and
continuity.
Within botany the individuals who paved the way for an ecological perspective
on the morphology and physiology of plants were Simon Schwendener (1829–1919)
and Gottlieb Haberlandt (1854–1945). In opposition to a science that – in response
to the speculative romantic biology of the early nineteenth century – focused exclu-
sively on “exact”, descriptive laboratory science and avoided speculation on the
function of specific histological or morphological structures in relation to the lives
of organisms, they founded what they called a “physiological anatomy”. The most
important hallmark of this research programme was the publication of Haberlandt’s
book Physiologische Pflanzenanatomie (Physiological plant anatomy) in 1884.63
Although not motivated by what we today would call “ecological questions”, the basic
62 Although see Höxtermann 2001 and, especially, the elaborate study by Cittadino (1990).
63 Haberlandt 1884. Five more (enlarged and revised) editions of the book were published before
1924. In 1919, an English translation (of the 4th edition) was published.
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19 Early Ecology in the German-Speaking World Through WWII
aim of this research programme was to identify the adaptive significance64 of plant
structures – involving the rigour of contemporary laboratory biology and even
experimental approaches, but with the need to leave the laboratory and examine
plants in their natural environments. Only in their natural environments, the researchers
felt, could the potential physiological usefulness of morphological studies be eluci-
dated (Schimper 1898, p. IV; Cittadino 1990). In order to see the potential differ-
ences in morphological adaptations to the environment more clearly (and supported
by the growing colonial ambitions of Germany) many biologists took the opportu-
nity to join expeditions headed for areas they perceived as extreme – or, at least from
a European perspective, exotic – environments, especially the tropics, but also the
deserts of Northern Africa. What began as a (comparative) analysis of the functional
significance of plant structures – guided by an evolutionary and also mostly
Darwinian perspective – gradually became a pillar for the ecology of plants in a
broader sense. In 1881 one of Schwendener’s first students in Berlin, Alexander
Tschirsch (1856–1939), wrote his PhD thesis on the relationship between the vegeta-
tion zones described by August Grisebach (1814–1879) and the specific anatomical
structures of the plants found in them,65 particularly those of the stomata of the
leaves. This was a first, important step in bringing together two hitherto independent
research fields: physiology and plant geography.
The idea was taken up by other botanists (e.g. Georg Volkens, 1855–1917,
another doctoral student of Schwendener) and culminated in Andreas Franz
Wilhelm Schimper’s book Pflanzengeographie auf physiologischer Grundlage
(Plant geography upon a physiological basis), which was published in 1898.
A.F.W. Schimper (1856–1901) was not directly related to the Schwendener circle,
but was a student of de Bary in Strasbourg, who was opposed to the ideas of
Schwendener and his collaborators. Only after his move to the University of Bonn
in 1882 and the experiences acquired from his travels to the tropics did he become
interested in ecological and biogeographical studies (Schenk 1901). His highly
influential book was based in part on his own research but was for the most part a
compilation of existing ecological knowledge relating to plant distribution and
adaptation. Approaching the subject matter from an evolutionary and selectionist
perspective (Cittadino 1990, p. 113), Schimper emphasized the need to examine the
causes of the different species of flora as the new goal of plant geography (Schimper
1898, p. III) and argued that ecology had to stay in close contact with experimental
plant physiology to serve this purpose (ibid., p. IV), furthering “ökologische
Pflanzengeographie” (ecological plant geography; ibid.).66 Schimper died only
64 Although the scientists in this school were united in the aim of looking at the evolutionary
significance of plant structures, some of the them (such as Schwendener himself, see Höxtermann
2001, p. 183) were opposed to Darwin’s theory of selection and adhered to a more Lamarckian
view of the mechanisms of evolution and adaptation.
65 These vegetation zones were based on the physiognomy of plants; see below.
66 Schimper made almost no reference to Eugenius Warming’s Plantesamfund, which was
published in 1895 (in Danish; German translation 1896, English translation 1909), whereas
Warming credited the German researcher’s work on physiological anatomy.
262
A. Schwarz and K. Jax
three years after the publication of his book – from a tropical disease contracted
during one of his journeys.
The tradition of physiological (or ecological) plant geography was influential –
not only in Germany but elsewhere too67 – and was also well received by zoologists
(e.g. Richard Hesse, see below). It did not, however, become the dominant tradition
in the new plant ecology, which was influenced to a much greater extent by the clas-
sificatory research programme developed in the tradition of Humboldt, Grisebach
and Drude and by the different schools of plant sociologists (see below). The most
notable follower of a physiological strand of plant geography was Heinrich Walter,68
who, however, remained an exception within German plant ecology.
The physiological programme, driven by an interest in validating Darwin’s
evolutionary theory, was also taken up by zoologists but, as in botany, this was
beyond the mainstream zoology of the time. The person who took Haeckel’s idea
of ecology as “external physiology” most seriously and tried to develop it into
a research programme was zoologist Carl Semper (1832–1893).69 Semper, a
morphologist at the University of Würzburg, was convinced that it was necessary
to collect more empirical evidence for Darwin’s theory, stating that “enough had
been done in the way of philosophising by the Darwinists, and the task that now lay
before us was to apply the test of exact investigations to the hypotheses we had
produced in this way”. (Semper 1881, p. v).70 This comment was also a sideswipe
at the speculative thinking of Haeckel. Thus Semper tried to combine the historical
and comparative approach of evolutionary theory and morphology with the causal
and even experimental research of the physiological biology of the time, both in
embryology, and – to a lesser degree – with respect to ecology (Nyhart 1995,
pp. 177ff.). As early as 1868 he postulated that it was necessary to investigate “the
influence of temperature, light, heat, humidity, nutrition, etc. on the living animal”
(die Einflüsse des Lichtes und der Wärme, des Feuchtigkeitsgrades, der Nahrung
etc. auf die lebenden Thiere) and to find “oecologische Gesetze”(ecological laws)
(Semper 1868, p. 229). The culmination of this area of his research came in a
lecture series which he held in 1877 at the Lowell Institute near Boston and which
was subsequently published in 1880 (German edition) and in 1881 (English edition).
This book, entitled Die natürlichen Existenzbedingungen der Thiere71 was the first
book on animal ecology ever to be published. While Semper still used Haeckel’s
terminology in his publication of 1868, there was no mention of “ecology” or
67 Thus, for example in the works of Frederic Clements and Henry Chandler Cowles; see Cittadino
1990, pp. 149ff. and Hagen 1988.
68 Walter’s students, especially Heinz Ellenberg and Wolfgang Haber, became highly influential in
post-WWII ecology in Germany.
69 Semper’s first name is spelt “Carl” in his early publications and “Karl” in later ones.
70 In the Geman version of 1880, the text reads thus: “...es sei von den Darwinisten doch schon
genug philosophirt, und die Aufgabe träte nun in ihr Recht, die auf diesem Wege gewonnenen
Hypothesen durch exacte Untersuchungen zu prüfen.” (Semper 1880, Volume 1, p. v)
71 Animal life as affected by the natural conditions of existence is the title of the American edition;
the title of the British edition is The conditions of natural existence as they affect animal life.
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19 Early Ecology in the German-Speaking World Through WWII
indeed any other reference to Haeckel’s wording in the later book.72 Making a similar
distinction to that made by Haeckel, Semper divides physiology into a “Physiologie
der Organe” (physiology of organs; i.e., physiology in the traditional sense) and a
“Physiologie der Organismen” (physiology of organisms), characterizing the latter
as “that branch of animal biology which regards the species of animals as actuali-
ties and investigates the reciprocal relations which adjust the balance between the
existence of any species and the natural, external conditions of its existence, in the
widest sense of the term.”73 His programme becomes explicit in the introduction to
the book, where he writes:
Da wir alle Theile des thierischen Körpers als echte Organe betrachten und die
Gesamtsumme ihrer Leistungen die Lebensfähigkeit der Arten bestimmen sehen, so erkennen
wir als Aufgabe des Zoologen zu untersuchen, wie die Lebensbedingungen auf die einzelnen
Thiere und ihre Organe wirken müssen, um zurück schließen zu können auf die physiolo-
gischen Ursachen des Entstehens verschiedener Thierformen.74
With Semper’s emphasis on explaining the morphological adaptations of animals to
their environment – in itself a means to furnish evidence of the causal mechanisms
of evolution – it comes as no surprise that Semper’s book is essentially (in modern
terminology) autecological: it deals with the individual organism (or the species
which it represents) and attempts to apply established laboratory methods to organisms
in their natural environment. Although his book is – rightly – credited with being
the first to point to food chains and trophic pyramids (concepts later formalized by
Victor Shelford and Charles Elton), this was done only in passing, namely in dealing
with food as one of the influences of the “inanimate” (sic!) environment. In fact, for
Semper, ecology served as a tool to explain morphology and evolution. If we look at
it differently, with the focus on ecology itself, we might justifiably say that Darwinian
evolutionary theory served here as the structuring idea for ecological research.
Although influential, Semper was not able to form any kind of continuing tradi-
tion or school in German ecology, nor was he able to carry out his programme of a
physiological animal geography himself. Animal ecology, as often acknowledged by
72 In a footnote to the ninth chapter, Semper (1880, Vol. 2, p. 268; 1881, p. 461) mentions Haeckel
as a major proponent of an extremly dogmatic Darwinism. As documented in his lecture Der
Haeckelismus in der Zoologie (Semper 1876), Semper had already developed an explicit distance
towards Haeckel and his work, which he (and others) perceived as overly speculative and therefore
unsound.
73 Semper 1881, p. 33. The German version (Semper 1880,Vol. 1, p. 39;) reads: “jenen Theil der
Biologie der Thiere […], welcher die Species der Thiere als Wirklichkeit ansieht und die
Beziehungen untersucht, welche zwischen der Existenz einer Art und ihren natürlichen äusseren
Existenzbedingungen obwalten (wobei dieser letztere Ausdruck natürlich in seinem weitesten
Sinne zu nehmen sein wird).”
74 Semper 1880, p. 28. The English version reads: “For since we consider all the parts of the
animal body as true organs, and see that the sum total of their functional activity determines
the vital fitness of the species, we perceive that it is the task of the zoologist to enquire how
the conditions of life must act upon individual animals and their organs, in order to be able to
deduce our inferences as to the physiological causes of the origin of different animal forms.”
(Semper 1881, p. 23).
264
A. Schwarz and K. Jax
its protagonists themselves, lagged behind plant ecology75 and, in particular, focused
more on species interactions and communities in Germany. Following Semper’s
seminal book it took until the 1920s for comprehensive overviews of animal ecology
to be produced in Germany. The tradition of physiological ecology and the merging
of physiology and biogeography was taken up most succinctly by Richard Hesse
(1868–1944). His book Tiergeographie auf ökologischer Grundlage (Animal
Geography upon an Ecological Basis) (1924) stands in the tradition of Semper and
of Schimper. As Hesse writes in his introduction, Schimper’s book was a shining
example for him, providing the basis on which he tried to model a zoological counter-
part text (Hesse 1924, p. V). Like these two scientists Hesse was an explicit Darwinst
and saw biogeography within the context of an adaptationist (and selectionist)
programme76: “Ecological animal geography considers the animals in their depen-
dence on the conditions of the area they live in, their ‘adaptedness’ to the conditions
of their environment, regardless of the geographical location of their living area”77;
and: “It is thus also one of the most important tasks of ecological animal geography
to investigate the ‘adaptations’ of the animals to their environment.”78
Like the physiological anatomist, he stresses the importance of experiments and
the need to extend biogeography beyond a purely descriptive method: “Everything
that has been called a process here is amenable to experimental confirmation and
physiological analysis.”79
Thus Hesse explicitly sees ecology in a physiological tradition: “These very
examples demonstrate how ecology is but a continuation and a complement of
physiological anatomy; the conditions of the environment are also included in the
intellectual linking up of the individual processes.”80 Without neglecting the role of
historical events in explaining biogeographical patterns, he expresses optimism
that ecology will be able to explain many phenomena by physical and chemical
75 E.g. Hesse 1924. p. 8: “Wir stehen zwar noch am Anfang einer experimentellen Ökologie;
besonders sind die Zoologen noch weit hinter den Botanikern zurück.” (We still are at the begin-
ning of an experimental ecology; in particular the zoologists are still far behind the botanists.);
Hesse 1927, p. 942: “Bis in die neueste Zeit wurden daher physiologische Anatomie und Ökologie
vernachlässigt, wenigstens von den Zoologen.” (Until very recently physiologogical anatomy and
ecology were neglected, at least by the zoologists).
76 See also Hartmann 1950 for Hesse’s position on Darwinism.
77 “Die ökologische Tiergeographie betrachtet die Tiere in ihrer Abhängigkeit von den Bedingungen
ihres Lebensgebietes, in ihrem ‘Angepaßtsein’ an ihre Umwelt, ohne Rücksicht auf die geogra-
phische Lage ihres Lebengebietes” (Hesse 1924, p. 6).
78 “So ist es auch eine der wichtigsten Aufgaben der ökologischen Tiergeographie die ‚Anpassungen’
der Tiere an ihre Umwelt zu untersuchen.” (ibid, p. 7).
79 “Alles, was hier als Vorgang bezeichnet wurde, ist einer experimentellen Bestätigung und physi-
ologischen Analyse zugänglich.” (Hesse 1924, p. 8).
80 “Gerade diese Beispiele zeigen, wie die Ökologie nur eine Fortführung und Ergänzung der
physiologischen Anatomie ist; es werden die Bedingungen der Umwelt mit einbezogen in die
gedankliche Verknüpfung der Einzelvorgänge.” (Hesse 1927, p. 944).
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19 Early Ecology in the German-Speaking World Through WWII
laws: “In animal ecology we thus encounter many events, which we can disassemble
into a sequence of processes. It is likely that it will be possible at some time to
reduce these processes to physical and chemical laws.”81
Hesse’s work was well received beyond the German–speaking world, and an
English translation of his 1924 book was published in 1937.82
The physiological and Darwinian approach never became a dominant tradition
within German ecology during the first half of the twentieth century, becoming less
important in comparison with the classificatory programme (i.e., plant sociology)
in plant ecology, the biocoenotic programme in animal ecology, and also the system
of lake types in aquatic ecology. A number of reasons may account for this decline
in importance. First, as a major driving force behind the programme, Darwinism
experienced what Julian Huxley called the “eclipse of Darwinism” (Bowler 1984),
marked especially by a great scepticism towards natural selection as the major
evolutionary mechanism. Many of the German ecologists who had been trained
during the time when anti-Darwinist critique was at its height among biologists, i.e.
between the 1880s and the 1930s, completely opposed the idea that the perceived
“harmony of nature” was the result of “mere chance”83 or at least ignored evolution-
ary issues. Especially in animal ecology, too, the importance of interactions of species
within communities (biocoenosis in German ecology) – or, from a functional point
of view, the roles of animal species within communities as well – seemed much more
conspicuous and important than the animal’s relations to abiotic factors, given the
greater mobility of most animal species compared with plants. This latter tradition of
German animal ecology is expressed most succinctly in the works of Karl August
Möbius, August Thienemann, and Karl Friederichs as discussed above.
The Classificatory Programme of Terrestrial Plant
Ecology: Plant Sociology
What we have called the classificatory programme here has its main roots in plant
geography. While biogeography also became important for the formation of ecology in
connection with the physiological approach, as shown above, another development
81 “So begegnen uns in der Ökologie der Tiere vielerlei Geschehnisse, die wir in eine Reihe von
Vorgängen auflösen können. Es wird voraussichtlich einmal gelingen, diese Vorgänge auf
physikalisch-chemische Gesetzmäßigkeiten zurückzuführen.” (Hesse 1927, p. 946).
82 Mitman (1992, p. 81) comments on the translation of Hesse’s book: “The volume was significant
because it made available in English one of the first books, apart from Shelford´s Animal
Communities in Temperate America, to offer an account of the worldwide distribution of animal
life on a physiological as opposed to a historical basis”.
83 E.g. Friederichs 1927, p. 156: “Es gibt die Einheit der Natur. Hätten wir mit diesem Bewußtsein
je im Banne der Darwinschen Theorie, die diese Einheit außer acht läßt, stehen können?” (The
unity of nature exists. Being aware of this, would we ever have been under the spell of Darwinian
theory, which disregards this unity?).
266
A. Schwarz and K. Jax
emphasized the distribution and environmental relations not of individual organisms
or species but of whole groups of species. It thus followed a route later labelled by
Schröter and Kirchner (1902) as “synecology”, namely: “the science of the plants
that live together, and at the same time the science of the plants that seek out analo-
gous ecological conditions.”84
The classificatory approach is rooted in Humboldtian plant geography, i.e. in the
early nineteenth century. Alexander von Humboldt (1769–1859) was the first to
describe systematically different recurring groups of plants. Humboldt’s classifica-
tion (Humboldt 1969), which he developed as a result of his travels to South
America, was based on physiognomic criteria, i.e. on plant form and not plant
taxonomy. The physiognomic approach was, however, for Humboldt not a purely
scientific one, but also explicitly related to aesthetic and emotional dimensions (see
Hard 1969; Kwa 2005). Humboldt’s ideas proved to be very influential. They gave
rise to several schools of plant geography which eventually, in a long process,
developed into an ecological plant geography, as one of the major pillars of ecology.85
In this process, Humboldt’s plant geography was gradually “cleansed” of its aesthetic
dimensions and enriched with concepts that today we call “ecological”. This
becomes especially evident with the development of the notion of plant formation,
being the first major concept of a unit describing whole assemblages of organisms
(here: plants). On the basis of Humboldt’s earlier classifications, August Grisebach
in 1838 defined the “formation” as an assemblage of plants composed of specific
“physiographic character”, i.e. based on plant forms.86
While Grisebach still referred to the aesthetic-emotional aspects of the physiognomic
perspective, his main aim was already to interpret the plant forms as an expression
of relations between vegetation and climate.87 Later authors, especially Grisebach’s
student Oscar Drude, teaching in Dresden (1890) and Danish botanist Eugenius
Warming (1895), completely abandoned Humboldt’s aesthetic dimensions. Drude
wrote: “It appeared necessary to me to remove the landscape-physiognomic aspects
as far as appropriate from the characteristics of the vegetation formations and
84 “Lehre von den Pflanzen, welche zusammen wohnen, und zugleich die Lehre von den Pflanzen,
welche analoge ökologische Bedingungen aufsuchen”. See also Chap. 14 this volume.
85 See Trepl 1987 and Nicolson 1996 for an extended account of this transition.
86 Grisebach 1838/1880, p. 2: “Ich möchte eine Gruppe von Pflanzen, die einen abgeschlossenen
physiographischen Charakter trägt, wie eine Wiese, ein Wald usw., eine pflanzengeographische
Formation nennen. Sie wird bald durch eine einzige gesellige Art, bald durch einen Complex von
vorherrschenden Arten derselben Familie charakterisirt, bald zeigt sie ein Aggregat von Arten,
die, mannigfaltig in ihrer Organisation, doch eine gemeinsame Eigenthümlichkeit haben, wie die
Alpentriften, die nur aus perennirenden Kräutern bestehen.” (I would term a group of terms which
bears a definite physiognomic character, such as meadow, a forest etc., a phytogeographic forma-
tion. The latter may be characterized by a single social species, by a complex of dominant species
belonging to one family, or finally, it may show an aggregate of species, which, though of various
taxonomic character, have a common pecularity; thus the alpine meadow consists almost entirely
of perennial herbs.) (translation as in Clements 1916, p. 116f.)
87 See also Trepl 1987 pp. 103–113 and Du Rietz 1931.
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19 Early Ecology in the German-Speaking World Through WWII
instead bring in the biological element”.88 And he adds: “Forest, shrubberies, and
meadows are different biological communities, which through their aggregation
prepare the natural habitat for similar plants or those dependent on them; that they
evoke a specific landscape impression is a highly enjoyable add-on, by which this
direction of botany becomes dear to the nature lover and valuable to the descriptive
geographer”.89
Bringing in the “biological element” here also meant that Drude brought in the
specific species (in contrast to just life forms) as crucial components of plant
communities, paving the way for the concept of plant association. Together with
Andreas Schimper’s (1898) systematic efforts to relate plant forms to abiotic condi-
tions and Warming’s (1895, 1896) emphasis on biological interactions between the
organisms of a plant community, a genuinely ecological plant geography (and thus
plant ecology) came into being and, with it, synecology in the modern sense.90
Drude, Warming and Schimper have often even been considered as the very founding
fathers of ecology as a discipline, (e.g. Worster 1985; Trepl 1987) because they
merged different strains of biological and geographical research, providing a new
perspective on the distribution of organisms, boosting what Allee et al. (1949) called
a “self-conscious discipline” of ecology. In any case, the works of these three
authors, all published in German, had a tremendous influence not only in the
German-speaking countries but far beyond. Thus the young Frederic Clements was
strongly influenced by the work of Drude (Tobey 1981), and Arthur Tansley also
refers to the books by Warming and Schimper as major inspirations for his work.91
Ecological plant geography and the ensuing synecological plant ecology was a
highly international enterprise from the beginning. Based on the seminal works of
Drude, Warming and Schimper, however, and in line with the different emphases of
their approaches, different schools of plant ecology developed very quickly.92 We will
88 Drude 1890, p. 23: “Es schien mir nämlich nötig, soweit als thunlich das landschaftlich-
physiognomische aus den Merkmalen der Vegetationsformationen zu entfernen und dafür das
biologische Element hineinzubringen”.
89 “Wälder, Gebüsche und Wiesen sind verschiedene biologische Gemeinden, welche durch ihren
Zusammenschluss ähnlich beanlagten oder auf sie angewiesenen Gewächsen die natürlichen
Standorte bereiten; dass sie einen bestimmten landschaftlichen Eindruck hervorrufen, ist eine
höchst angenehme Zugabe, durch welche diese Richtung der Botanik dem Naturfreunde lieb, dem
beschreibenden Geographen wertvoll wird.” (ibd.)
90 Schimper’s work (especially Schimper 1898) is located at the boundary between the physiologi-
cal approach and the synecological one.
91 The original edition of Warming’s book Plantesamfund was published in Danish in 1895. Most
readers outside Denmark, however, used the German version of the book of 1896 (see e.g. Tansley
1947). A (strongly modified) English version of Planetsamfund appeared, somewhat tardily, in
1909. On Warming and his seminal book see Anker Chap. 23, this volume.
92 Overviews of the different schools of vegetation science can be found in Whittaker 1962,
Mueller-Dombois & Ellenberg 1974; Shimwell 1971; Dierschke 1994; for the Russian tradition in
particular and its politically hastened decline since the mid-1930s, see Weiner 1984, 1988. The
early history of the discipline and its theory, in which the characterization of ecological units plays
a major role, is described in Clements 1916; Rübel 1917, 1920; Gams 1918; and Du Rietz 1921.
268
A. Schwarz and K. Jax
describe here in particular those relating to the German-speaking world, which
focused on an approach that came to be known as plant sociology.
The young science of synecological plant ecology (or vegetation ecology) saw
heated debates early on, both regarding the definition and “nature” of the basic
units of the discipline (in particular plant formation and plant association) and the
appropriate methods to describe and classify these units and their dynamics. As a
result of divergent opinions regarding the meaning of pivotal terms and the contents
of the concepts connected with them, a report and accompanying proposal for the
nomenclature of phytogeography (Flahault and Schröter 1910) was elaborated for
the III. International Congress of Botany in Brussels. The aim of this proposal was
to arrive at a uniform usage of the words “formation”, “association” and others.
This attempt was initially unsuccessful, however. In addition, a deeper split regarding
the view of plant assemblages became apparent around the beginning of the twen-
tieth century, which led to divided traditions of vegetation ecology in continental
Europe and the Anglo-Saxon countries, a rift that can be perceived even today. On
the one side there was the mostly descriptive and classificatory plant sociology in
the tradition of Schröter, Sernander and later especially Braun-Blanquet, Du Rietz,
and Tüxen, and on the other the more dynamically oriented vegetation ecology, i.e.
oriented towards the notion of “development” and succession, as shaped by
Clements, Tansley and Gleason, with a common theoretical basis but also with clear
and specific differences.93
The main schools of European plant sociology here were the Central European
“Zurich-Montpellier-school”, founded by Carl Schröter and Charles Flahault, and
the Scandinavian “Upsala school”, founded by Rutger Sernander. These designations
were not strict geographic ones: Austrian botanist Helmut Gams – although an
exception – was considered part of the Upsala school, for example. Russia also had –
especially up until the 1930s – a theoretically and empirically important tradition
of research in plant sociology. The competing schools of plant sociology differed
in many theoretical and methodological points, over which arguments were
exchanged with great fervour; these disputes were, as Whittaker (1962, p. 27) put
it, “waged with the special intensity of a civil war”. Differences concerned things
as the appropriate spatial dimension of their major ecological unit, namely the plant
association, questions of quantitative vs. qualitative methods for its characterization,
or the questions which kinds of species should be selected to characterize the associa-
tion: constants, as species occurring in 90% or more of the samples, or character
species, as species with a narrow ecological amplitude and thus restricted to particlar
associations. A more fundamental difference concerned the question if associations
only have to be “found” in nature or if are they a mere product of the ordering
human mind. Is the association a concrete thing or a class concept? Du Rietz, for
93 The British ecologists (as represented by the British Vegetation Committee, the predecessor of
the British Ecological Society) included a whole successional sere into their definitions and clas-
sifications of plant formation, which most central European ecologists rejected as too full of
hypotheses (see Flahault and Schröter 1910).
269
19 Early Ecology in the German-Speaking World Through WWII
example insisted on the “reality” of the association: “The associations like the spe-
cies are not produced in scientific treatments and textbooks. They are species com-
binations existing in nature and delimited more or less sharply by nature itself.”94
In contrast, Braun-Blanquet in the same year (1921) emphasized: “There is, by
and large, agreement that the association as well as the species is an abstraction,
while we are faced with single association individuals and local stands in nature.”95
This question, about the “reality” of the plant association, evoked a flood of
controversial publications, and Du Rietz called the issue in 1928 “one of the most
important and burning main problems of modern plant sociology” (eine[s] der
wichtigsten und aktuellsten Hauptprobleme der modernen Pflanzensoziologie;
Du Rietz 1928, p. 20).96
Although there had always been publications in each native language (in particular
French, Danish, Swedish, Norwegian and Russian), the main working language in
continental European plant sociology was German. Plant sociology was an explicit
research programme and became for a long time the dominant research tradition of
plant ecology in the German-speaking world.
Josias Braun-Blanquet, a student of Flahault and Schröter and soon the leading
figure in the Zurich-Montepellier school, developed the most detailed and most
influential research programme for plant sociology, laid out in detail in his book
Pflanzensoziologie (Plant Sociology) of 1928.97 In the first pages of this book,
Braun-Blanquet describes the status and overall aims of the field. First of all, “plant
sociology” is considered to be synonymous with “vegetation science”, which he, like
many others (e.g. Du Rietz 1921) perceives as a distinct discipline, and not just as a
sub-discipline of ecology or geography (Braun-Blanquet 1928, p. III). The object of
the discipline is the “plant community as a social unit”98 and its “clear but remote
aim”99 is “the characterisation and description of the social units, their causal explana-
tion, the study of their development and distribution and their clear and systematic
arrangement”.100 The research field is then characterized (Braun-Blanquet 1928, p. 1f.)
by five “main problems”, relating to (1) the organization and structure of plant
societies, (2) their ecological relations (3) their (successional) development, (4) their
spatial distribution and (5) their classification and systematics.
94 Du Rietz 1921, p. 15; emphasis in original. “Die Assoziationen ebenso wie die Arten werden nicht
in wissenschaftlichen Abhandlungen und Lehrbüchern fabriziert. Sie sind in der Natur existierende,
durch die Natur selbst mehr oder minder scharf und deutlich abgegrenzte Artenkombinationen”.
95 Braun-Blanquet 1921, p. 311. “Man ist heute im grossen ganzen darüber einig, dass die
Assoziation so gut wie die Art eine Abstraktion darstellt, während uns in der Natur einzelne
Assoziationsindividuen oder Lokalbestände entgegentreten.”
96 See Jax 2002, pp. 110ff. for an more detailed analysis of this controversy.
97 English translation 1932.
98 “Pflanzengesellschaft als soziale Einheit”; ibid.
99 “klares aber fernes Ziel”; ibid.
100
“die Fassung und Beschreibung der Gesellschaftseinheiten, ihre kausale Erklärung, das Studium
ihrer Entwicklung und Verbreitung und ihre übersichtliche systematische Anordnung”; ibid.
270
A. Schwarz and K. Jax
In a similar manner, Du Rietz, a student of Sernander and the dominant exponent
of the Uppsala school, explained some years earlier what he considered to be the
“Endziel” (ultimate aim) of plant sociology, namely:
“eine[r] allseitige[n] Kenntnis von den in der Natur existierenden Pflanzengesellschaften,
ihrem Aussehen, ihrer Zusammensetzung und ihrem inneren Bau, ihrer Entstehung und
ihren Veränderungen, ihrer Verbreitung und Verteilung auf der Erde, ihren
Lebensverhältnissen und ihrer Sukzession, nicht aber darin, daß man einzelne von diesen
vielseitigen Forschungsaufgaben auf ein Piedestal über alle übrigen erhebt.”101
In spite of many theoretical and methodological differences between the two schools
of plant sociology, the overall research programme was thus very similar, and equally
broad and ambitious. The direction and actual practice of research during the following
decades, however, was much narrower, which already was anticipated by both Braun-
Blanquet and Du Rietz. They both saw the description and assessment of plant societies
as the first (although transient) task of plant sociology.102 Description and classification
was in fact the major work done within plant sociology in the German-speaking coun-
tries up until the end of WWII.103 It was ecological in the sense of Drude’s and
Schimper’s work, as it laid great stress on elucidating the relationships between plant
communities and their (abiotic) environment. What was largely lacking, however, was
an investigation of the species’ relationships within the communities, as emphasized
by Warming. The actual classificatory programme of terrestrial plant ecology in effect
provided a huge amount of valuable empirical and spatially concrete data on the dis-
tribution of plant communities and their relation to factors such as climate and soil,
as well as new methodologies. However, it did not produce much theoretical progress
in the explication of plant communities and their dynamics.
Summary
To draw this overview to an end, let us briefly recapitulate some of the issues that
have arisen in our account of German-speaking ecology. German-speaking ecology
develops in two strands from the beginning – aquatic and terrestrial ecology. These
evolve at different rates. This becomes apparent with respect to the institutional,
cognitive and epistemic context. In aquatic ecology a process of institutionalization
101 Du Rietz 1921, p. 248: “a general knowledge of the plant societies existing in nature, their
appearance, their composition and their internal structure, their origin and their changes, their distribu-
tion and arrangement on the earth, their living conditions and their succession, but [its aim is] not
to raise up any single one of these various research tasks on a pedestal above all the others”.
This text appears towards the very end of Du Rietz’s PhD thesis about the methodological founda-
tion of modern plant sociology” (Zur methodologischen Grundlage der modernen Pflanzensoziologie).
102 Du Rietz (ibid.), referring to his statement above, said that if one aspect were to gain some
prominence, then it would have to be “the determination of the plant societies existing in nature
and of their natural boundaries” (die Feststellung der in der Natur existierenden Pflanzengesellschaf-
ten und ihrer natürlichen Grenzen) – as a necessary prerequiste for all the other tasks.
103 See e.g. Dierschke 1994, p. 20.
271
19 Early Ecology in the German-Speaking World Through WWII
takes place between the 1890s and the 1920s: laboratories and field stations, posi-
tions, scientific societies and journals are all established. Terrestrial ecology – espe-
cially plant ecology and plant geography – also develops these structures, yet remains a
rather disparate field, especially when it comes to animal ecology, until about the
1920s. We have attempted to take account of these differences by proposing two
different conceptual patterns to accommodate the different structures and transfor-
mations in the respective fields of knowledge. Of course, we are aware that this is
just an initial historical reconstruction, which still contains many unrelated frag-
ments and cognititve fissures. Despite this, we are convinced that it is time to rear-
range at least some of the narratives about German-speaking ecology while adjusting
the vantage point on hitherto unseen ruptures and associations, most of them due to
these differences between aquatic and terrestrial ecology. Taking them into consid-
eration will hopefully facilitate the development of a novel and productive perspec-
tive on the constitution of ecological knowledge in the German-speaking world.
References
Allee WC, Emerson AE, Park O, Park T, Schmidt KP (1949) Principles of animal ecology.
Saunders, Philadelphia
Bowler PJ (1984) Evolution. The history of an idea. University of California Press, Berkeley
Braun-Blanquet J (1921) Prinzipien einer Systematik der Pflanzengesellschaften auf floristischer
Grundlage. Jahrbuch Sankt Gallener Naturwissenschaftlichen Ges 57:305–351
Braun-Blanquet J (1928) Pflanzensoziologie. Springer, Berlin
Brehm V (1930) Einführung in die Limnologie. Springer, Berlin
Burckhardt G (1900) Quantitative Studien über das Zooplankton des Vierwaldstättersees. Mitt
Naturforschenden Ges Luzern 3:129–411, 686–707, 414–434
Burkamp W (1929) Die Struktur der Ganzheiten. Junker und Dünnhaupt, Berlin
Busch B (ed) (2007) Jetzt ist die Landschaft ein Katalog voller Wörter. Beiträge zur Sprache der
Ökologie. Valerio 5, Die Heftreihe der Deutschen Akademie für Sprache und Dichtung.
Göttingen, Wallstein
Cittadino E (1990) Nature as the laboratory. Darwinian plant ecology in the German Empire,
1880–1900. Cambridge University Press, Cambridge
Clements FE (1916) Plant succession. An analysis of the development of vegetation. Carnegie
Institution of Washington, Washington, DC, Publication No. 242
Dahl F (1921) Grundlagen einer ökologischen Tiergeographie. Gustav Fischer, Jena
Daum AW (1998) Wissenschaftspopularisierung im 19. Jahrhundert. Bürgerliche Kultur, natur-
wissenschaftliche Bildung und die deutsche Öffentlichkeit, 1848–1914. Oldenbourg Verlag,
München
Dierschke H (1994) Pflanzensoziologie. Ulmer, Stuttgart
Drude O (1890) Handbuch der Pflanzengeographie. Verlag von J. Engelhorn, Stuttgart
Du Rietz GE (1921) Zur methodologischen Grundlage der modernen Pflanzensoziologie. Adolf
Holzhausen, Wien
Du Rietz GE (1928) Kritik an pflanzensoziologischen Kritikern. Botaniska Notiser 1–30
Du Rietz GE (1931) Life-forms of terrestrial flowering plants. Acta Phytogeographica Suecica III:1–95
Flahault C, Schröter C (eds) (1910) Phytogeographische Nomenklatur. III. Internationaler
Botanischer Kongress, Brüssel 1910. Zürcher & Furrer, Zürich
Forel F-A (1886) Programme d’études limnologiques pour les lacs subalpins. Arch Sci Phys Nat
3:548–550
Forel F-A (1892–1904) Le Léman. Monographie limnologique, vol 1–3. F. Rouge, Lausanne
272
A. Schwarz and K. Jax
Forel F-A (1901) Handbuch der Seenkunde. Allgemeine Limnologie. Engelhorn, Stuttgart
Forel FA (1896) La limnologie, branche de la géographie. Rep. Sixth Int. Geogr. Congress held in
London 1895 593–602
Frič A, Václav V (1894) Untersuchungen über die Fauna der Gewässer Böhmens IV. Die
Thierwelt des Unterpočernitzer und Gatterschlager Teiches. Archiv für die naturwissen-
schaftliche Landesdurchforschung von Böhmen 9, Prag
Friederichs K (1927) Grundsätzliches über die Lebenseinheiten höherer Ordnung und den ökolo-
gischen Einheitsfaktor. Naturwissenschaften 8:153–157, 182–186
Friederichs K (1930) Die Grundfragen und Gesetzmäßigkeiten der land- und forstwirtschaftli-
chen Zoologie (insbesondere der Entomologie), vol 1, 2. Verlagsbuchhandlung Paul Parey,
Berlin
Gams H (1918) Prinzipienfragen der Vegetationsforschung. Ein Beitrag zur Begriffsklärung und
Methodik der Biocoenologie. Vierteljahresschridft der Naturforschenden Gesellschaft Zürich
63:293–493
Grisebach A (1838) Über den Einfluß des Klimas auf die Begrenzung der natürlichen Floren. In:
Grisebach A (ed) Gesammelte Abhandlungen und kleinere Schriften zur Pflanzengeographie.
Verlag von Wilhelm Engelmann, Leipzig, pp 1–29
Haberlandt G (1884) Physiologische Pflanzenanatomie. Engelmann, Leipzig
Hagen JB (1988) Organism and environment. Frederic Clements´s vision of a unified physiologi-
cal ecology. In: Rainger R, Benson KR, Maienschein J (eds) The American development of
biology. University of Pennsylvania Press, Philadelphia, pp 257–280
Hard G (1969) “Kosmos” und “Landschaft”. Kosmologische und landschaftsphysiognomische
Denkmotive bei Alexander von Humboldt und in der geographischen Humboldt-Auslegung
des 20. Jahrhunderts. In: Pfeiffer H (ed) Alexander von Humboldt. Werk und Weltgeltung.
Piper-Verlag, München, pp 133–177
Hartmann M (1950) Nachruf auf Richard Hesse. - Jahrbuch der Deutschen Akademie der
Wissenschaften zu Berlin, 1946–1949, pp 160–170
Hentschel E (1909) Das Leben des Süßwassers. Eine gemeinverständliche Biologie. Ernst
Reinhardt, München
Hentschel E (1923) Grundzüge der Hydrobiologie. Gustav Fischer, Jena
Hesse R (1924) Tiergeographie auf ökologischer Grundlage. Gustav Fischer, Jena
Hesse R (1927) Die Ökologie der Tiere, ihre Wege und Ziele. Naturwissenschaften 15:942–946
Höxtermann E (2001) Die Schwendener-Schule der Physiologischen Anatomie - ein “Grundpfeiler”
der Pflanzenökologie. Verhandlungen zur Geschichte und Theorie der Biologie 7:165–189
Jax K (1998) Holocoen and ecosystem. On the origin and historical consequences of two concepts.
J Hist Biol 31:113–142
Jax K (2002) Die Einheiten der Ökologie. Analyse, Methodenentwicklung und Anwendung in
Ökologie und Naturschutz. Peter Lang, Frankfurt
Junge F (1885) Der Dorfteich als Lebensgemeinschaft. Lipsius & Tischer, Kiel
Karny HH (1934) Biologie der Wasserinsekten. Ein Lehr- und Nachschlagewerk über die wich-
tigsten Ergebnisse der Hydro-Entomologie. Fritz Walter, Wien
Kluge T, Schramm E (1986) Wassernöte. Sozial- und Umweltgeschichte des Trinkwassers. Alano-
Verlag, Aachen
Kohler RE (2002) Landscapes and labscapes: Exploring the lab-field frontier in biology. The
University of Chicago Press, Chicago
Kretschmann C (2006) Räume öffnen sich. Naturhistorische Museen im Deutschland des 19.
Jahrhunderts. Akademie-Verlag, Berlin
Kuhn TS (1988) Die Struktur wissenschaftlicher Revolutionen. Suhrkamp, Frankfurt am Main
Kwa C (2005) Alexander von Humboldt’s invention of the natural landscape. Eur Legacy
10:149–162
Lenz F (1928) Einführung in die Biologie der Süsswasserseen. Biologische Studienbücher, vol IX.
Berlin, Julius Springer
Mitman G (1992) The state of nature. Ecology, community, and American social thought,
1900–1950. University of Chicago Press, Chicago
273
19 Early Ecology in the German-Speaking World Through WWII
Möbius KA (1883) The oyster and oyster culture. Report of the comissioner for 1880. United States
Comission of Fish and Fisheries. Government Printing Office, Washington, DC, pp 683–751
Möbius KA (2006) Zum Biozönose-Begriff. Die Auster und die Austernwirtschaft 1877 (2nd ed.
by Thomas Potthast; 1st edition and comment by Günther Leps 1986). – Frankfurt am Main:
Harri Deutsch
Mueller-Dombois D, Ellenberg H (1974) Aims and methods of vegetation ecology. Wiley, New York
Müller-Navarra S (2005) Ein vergessenes Kapitel der Seenforschung. Martin Meidenbauer
Verlagsbuchhandlung, München
Naumann E (1918a) Försök angående vissa avfallsprodukters och gödselämnens inverkan på vattnets
biologi. Särtryck Ur Skrifter, Utgivna Av Södra Sveriges Fiskeriförening 1917 (3–4):10–44
Naumann E (1918b) Undersökningar över fytoplanktonproduktionen i dammar vid aneboda 1917.
Sartryck Ur Skrifter, Utgivna Av Södra Sveriges Fiskeriförening 1:62–75
Naumann E (1921) Einige Grundlinien der regionalen Limnologie. Lunds Univesitets Årsskrift
NF 17:1–22
Nicolson M (1996) Humboldtian plant geography after Humboldt: the link to ecology. Br J Hist
Sci 29:289–310
Nyhart LK (1995) Biology takes form. Animal morphology and the German universities, 1800 –
1900. University of Chicago Press, Chicago
Pörksen U (2001) Was spricht dafür das Deutsche als Naturwissenschaftssprache zu erhalten?
Abhandlungen der Deutschen Akademie der Naturforscher Leopoldina NF 87:5–31
Potthast T (2003) Wissenschaftliche Ökologie und Naturschutz: Szenen einer Annäherung. In: Radkau J,
Uekötter F (eds) Naturschutz und Nationalsozialismus. Campus, Frankfurt, pp 225–256
Psenner R, Alfreider A, Schwarz AE (2008) Aquatic microbial ecology: water desert, microcosm,
ecosystem. What comes next? Int Rev Hydrobiol 93:606–623
Radkau J (2003) Naturschutz und Nationalsozialismus – wo ist das Problem? In: Radkau J,
Uekötter F (eds) Naturschutz und Nationalsozialismus. Campus, Frankfurt, pp 41–55
RegioWasser eV (ed) (2009) 50 Jahre Rheinforschung. Lebensgang und Schaffen eines deutschen
Naturforschers Robert Lauterborn (1869–1952). Lavori Verlag, Freiburg
Rübel E (1917) Anfänge und Ziele der Geobotanik. Vierteljahresschrift der Naturforschenden
Gesellschaft Zürich 62:629–650
Rübel E (1920) Die Entwicklung der Pflanzensoziologie. Vierteljahresschrift der Naturforschenden
Gesellschaft Zürich 65:573–604
Schenk H (1901) A.F. Wilhelm Schimper. Berichte der Deutschen Botanischen Gesellschaft
19:954–970
Schimper AFW (1898) Pflanzengeographie auf physiologischer Grundlage. Gustav Fischer, Jena
Schröter C, Kirchner O (1896) Vegetation des Bodensees. 1. Band. Stettner, Lindau
Schröter C, Kirchner O (1902) Die Vegetation des Bodensees, 2. Teil. - Lindau: Kommissionsverlag
der Schriften des Vereins der Geschichte des Bodensees und seiner Umgebung von Joh.
Stettner, Thom
Schwabe GH (1961) August Thienemann in memoriam. Oikos 12:310–316
Schwarz AE (2003a) Wasserwüste - Mikrokosmos - Ökosystem. Eine Geschichte der Eroberung
des Wasserraumes. Rombach-Verlag, Freiburg
Schwarz AE (2003b) Die Ökologie des Sees. Diagramme als Theoriebilder. Bildwelten des
Wissens Kunsthistorisches Jahrbuch für Bildkritik 1:64–74
Semper K (1868) Reisen im Archipel der Phillipinen. Zweiter Teil: Wissenschaftliche Resultate.
Erster Band: Holothurien. Verlag von Wilhelm Engelmann, Leipzig
Semper K (1876) Der Haeckelismus in der Zoologie. W. Maukes Söhne, Hamburg
Semper K (1880) Die natürlichen Existenzbedingungen der Thiere. Brockhaus, Leipzig
Semper K (1881) Animal life as affected by the n-atural conditions of existence. D. Appleton &
Co, New York
Shimwell DW (1971) The description and classification of vegetation. University of Washington
Press, Seattle
Steinecke F (1940) Der Süßwassersee. Die Lebensgemeinschaften des nährstoffreichen Binnensees.
Quelle und Meyer, Leipzig
274
A. Schwarz and K. Jax
Steleanu A (1989) Geschichte der Limnologie und ihrer Grundlagen. Haag und Herchen,
Frankfurt am Main
Tansley AG (1947) The early history of modern plant ecology in Britain. J Ecol 35:130–137
Thienemann A (1921) Seetypen. Naturwissenschaften 9:343–346
Thienemann A (1923) Zwecke und Ziele der Internationalen Vereinigung für theoretische und
angewandte Limnologie. Verhandlungen der Internationalen Vereinigung für theoretische und
angewandte Limnologie 1:1–5
Thienemann A (1925) Der See als Lebenseinheit. Naturwissenschaften 13:489–600
Thienemann A (1926) Limnologie. Eine Einführung in die biologischen Probleme der
Süßwasserforschung. - Breslau
Thienemann A (1927) Der Nahrungskreislauf im Wasser. 31. Jahresversammlung zu Kiel 1926.
Zoologischer Anzeiger (Verhandlungen der Deutschen Zoologischen Gesellschaft 31)
2(Supplementband):29–79
Thienemann A (1933) Vom Wesen der Limnologie und ihrer Bedeutung für die Kultur der
Gegenwart. Verhandlungen der Internationalen Vereinigung für theoretische und angewandte
Limnologie 6:21–30
Thienemann A (1935) Die Bedeutung der Limnologie für die Kultur der Gegenwart.
Schweizerbart’sche Verlagsbuchhandlung, Stuttgart
Thienemann A (1939) Grundzüge einer allgemeinen Ökologie. Schweizerbart’sche
Verlagsbuchhandlung, Stuttgart
Thienemann A (1942) Vom Wesen der Ökologie. - Biologia Generalis 3/4 (special
edition):312–331
Thienemann A (1951) Vom Gebrauch und vom Mißbrauch der Gewässer in einem Kulturlande.
Arch Hydrobiol 45:557–583
Thienemann A (1954) Wasser - Das Blut der Erde. In: Uns ruft der Wald. Handbuch der
Schutzgemeinschaft Deutscher Wald. Rheinhausen: Verlagsanstalt Rheinhausen, pp 45–49
Tobey RC (1981) Saving the prairies. The life cycles of the founding school of American plant
ecology, 1895–1955. University of California Press, Berkeley
Trepl L (1987) Geschichte der Ökologie. Vom 17. Jahrhundert bis zur Gegenwart. Athenäum,
Frankfurt am Main
Tümmers HJ (1999) Der Rhein. Ein europäischer Fluss und seine Geschichte. Beck, München
Ule W (1901) Der Würmsee (Starnbergersee) in Oberbayern, eine limnologische Studie. Leipzig
Ule W (1906) Studien am Ammersee in Oberbayern. Riedel, München
Utermöhl H (1925) Limnologische Phytoplanktonstudien: Die Besiedelung ostholsteinischer Seen
mit Schwebpflanzen. - Archiv für Hydrobiologie, Suppl. 5
von Humboldt A (1969) In: Meyer-Abich A (ed) Ansichten der Natur. Reclam, Stuttgart
Walter H (1927) Einführung in die allgemeine Pflanzengeographie Deutschlands. Gustav Fischer,
Jena
Ward HB (1899) The freshwater biological stations of the world. Science 9:497–507
Warming E (1895) Plantesamfund. Grundtræk af den økologiske plantegeografi S. Philipsen,
Kjobenhavn
Warming E (1896) Lehrbuch der ökologischen Pflanzengeographie. Eine Einführung in die
Kenntnis der Pflanzenvereine. Gebrüder Bornträger, Berlin
Warming E (1909) Oecology of plants: An introduction to the study of plant communities. Oxford
University Press, Oxford
Wasmund E (1926) Wissenschaftsprovinzen. Deutsche Rundschau 52(12):243–253
Weiner DR (1984) Community ecology in Stalin’s Russia: “Socialist and bourgeois” science. Isis
75:684–696
Weiner DR (1988) Ecology, conservation, and cultural revolution in Soviet Russia. Indiana
University Press, Bloomington & Indianapolis
Weismann A (1877) Das Thierleben im Bodensee. Schriften des Vereins für Geschichte des
Bodensees und seiner Umgebung 7:132–161
Whittaker RH (1962) Classification of natural communities. Bot Rev 28:1–239
275
Woltereck R (1928) Über die Spezifität des Lebensraumes, der Nahrung und der Körperformen
bei pelagischen Cladoceren und über “Ökologische Gestalt-Systeme”. Biologisches Zentralblatt
48:521–551
Woltereck R (1940) Ontologie des Lebendigen. Ferdinand Enke, Stuttgart
Worster D (1985) Nature’s economy. A history of ecological ideas. Cambridge University Press,
Cambridge
Zacharias O (1888) Vorschlag zur Gründung von zoologischen Stationen behufs Beobachtung der
Süßwasser-Fauna. Zool Anz 11:18–27
Zacharias O (1904) Skizze eines Spezial-Programms für Fischereiwissenschaftliche Forschungen.
Fischerei-Zeitung 7:112–115
Zacharias O (1905) Über die systematische Durchforschung der Binnengewässer und ihre
Beziehung zu den Aufgaben der allgemeinen Wissenschaft vom Leben. Forschungsberichte
aus der biologischen Station Plön 12:1–39
Zacharias O (1907) Das Süßwasserplankton. Teubner, Leipzig
19 Early Ecology in the German-Speaking World Through WWII
277
Chapter 20
The History of Early British
and US-American Ecology to 1950
Robert McIntosh
Introduction
Scientific ecology was anticipated by a long history of natural history observations
from classical times, recently termed protoecology (Glacken 1967; Egerton 1976).
Ecology (Oekologie) was coined in 1866 by Ernst Haeckel, a German biologist and
advocate of Darwin (see Chap. 10). One of the earliest uses in English was by
Patrick Geddes, a British botanist. In 1880, 20 years before the term ecology was
in general use, Geddes offered a hierarchy of the sciences putting ecology under
sociology rather than biology (Mairet 1957), anticipating later connections with
sociology. Geddes taught the brothers Robert and William Smith who later joined
with Arthur G. Tansley in furthering vegetation studies and plant ecology in Britain
(Tansley 1911). In 1893 the president of the British Association for the Advancement
of Science described “oecology” as a branch of biology coequal with morphology
and physiology and “by far the most attractive” (McIntosh 1985).
Tansley was influential in establishing a British Vegetation Committee in 1904
and in producing a volume on Types of British Vegetation in 1911 and organizing a
field trip, which marked the first meeting of the pioneer British and American plant
ecologists with continental ecologists. The British Ecological Society and the
Journal of Ecology were initiated in 1913 and Tansley gave the first presidential
address to any ecological society in 1914.
In the United States formalization of ecology similarly lagged behind coinage of
the term. Among its earliest proponents was Stephen Alfred Forbes. Following
service in the Civil War Forbes undertook extensive studies in natural history
including insects, fish and birds, initiating an Illinois State Laboratory of Natural
History in 1877. Forbes produced many of the earliest and most insightful publica-
tions on ecology notably, in 1887, The Lake as a Microcosm, treating the lake as
R. McIntosh (*)
Formerly Professor at the Department of Biological Sciences, University of Notre Dame,
Notre Dame, Indiana, USA
A. Schwarz and K. Jax (eds.), Ecology Revisited: Reflecting on Concepts, Advancing Science,
DOI 10.1007/978-90-481-9744-6_20, © Springer Science+Business Media B.V. 2011
278
R. McIntosh
producing an equilibrium, stressing its holistic nature; and in 1894 he recognized
ecology as a science including “economic entomology”, “the whole Darwinian
doctrine” and agriculture (Croker 2001).
Early ecology in the United States flourished largely in the Midwest, in universi-
ties and state natural history agencies. In Wisconsin E.A. Birge initiated limnologi-
cal studies of plankton in the 1890s as director of the State Natural History Division,
continuing these and other aquatic studies with Chancey Juday for three decades at
the University of Wisconsin. Plant ecology in the US, as in Britain, was prominent
in early ecology, notably at the universities of Nebraska, Minnesota and Chicago. In
1893 the Madison Botanical Congress formally adopted the term ecology, dropping
the earlier diphthong “oe”. Prominent among early plant ecologists were F.E.
Clements at the Universities of Nebraska and Minnesota and H.C. Cowles of the
University of Chicago. Clements became the major systematizer for plant ecology
and produced two of its seminal volumes (Clements 1905, 1916). Clements consid-
ered “dynamic ecology”, emphasizing the succession of the plant community to a
stable endpoint, the climax association, under the control of the climate. Clements
regarded the community as an integrated organism or even a superorganism. (see
Chap. 4) In 1916 he formulated his ideas as a “universal law” and organismic con-
cepts constituted the major synthesis of early twentieth century ecology.
The concept of nature as an organism, or superorganism, grew out of the tradi-
tion of a divinely organized balance of nature and was metaphorically extended to
the complex of organisms or community. S.A. Forbes wrote, “A group or associa-
tion of animals or plants is like a single organism” Clements, and his co-author
Victor Shelford, stated that the organismic concept in ecology “is a veritable magna
carta for future progress” (Clements and Shelford 1939). F.S. Bodenheimer wrote
“every modern textbook of ecology stresses the highly integrated supraorganismic
structure of communities” but noted, “there is no scientific evidence to support it”,
an observation commonly ignored (McIntosh 1998).
Cowles, working on the sand dunes of the Lake Michigan shore, based his ideas,
like Clements, on succession stemming from studies of vegetation. He differed
from Clements in recognizing succession as a tortuous process not leading to a
stable climax. His famous axiom was that succession is “a variable approaching a
variable rather than a constant (…)”, a problem frequently faced by ecologists.
Animal ecology followed on the heels of plant ecology in the US and Britain. In
the US animal ecology was advanced by C.C. Adams and Victor Shelford, both
having early association with S.A. Forbes in Illinois. Adams and Shelford contrib-
uted to the study of animal communities and provided important early volumes on
animal ecology (Adams 1913; Shelford 1913). Adams (1935) was an early observer
of the linkage of general ecology to human ecology. Both were involved in the
formation of the Ecological Society of America in 1914, and Victor Shelford was
its first president. Its journal, Ecology, was founded in 1920.
British animal ecology was stimulated by Charles Elton’s (1927) primer, Animal
Ecology, following his extended surveys (1921–1924) of Arctic animal communi-
ties. Elton elaborated key ideas of community organization, food chain (trophic
structure) pyramid of numbers, niche and, like Charles Adams, described ecology
279
20 The History of Early British and US-American Ecology to 1950
as scientific natural history. Elton’s long association with the Hudson’s Bay
Company and its fur data provided early insights into population dynamics of
predator and prey. Elton established a research area, Wytham Woods, in Oxfordshire
that became one of the most studied areas on earth (Cox 1979) and was later used
as a site for one of the well known Inspector Morse mystery series on TV.
Populations and Mathematics
Ecology developed as a loose amalgam of marine biology, limnology and plant and
animal ecology. Oddly, parasitology, an intrinsically ecological study, remained
largely separate from ecology until the 1960s. Population ecology developed as a
major element of ecology as the study of the numbers of individuals of a species,
the changes in numbers and interactions among species, such as predation of birds
on insects, a one-time concern of Benjamin Franklin’s, and competition between
species. Human population growth described by Thomas Malthus (1798) was a
major stimulus to Charles Darwin’s theory of evolution.
Population counts, or censuses, were a common aspect of early ecology, fol-
lowed by the rise of statistics in ecology. In 1928 Aldo Leopold began his studies
of game populations in the US that led to his pioneer volume on Game Management
and subsequent career in ecology at the University of Wisconsin. Charles Elton, in
Britain, formed a Bureau of Animal Populations and the Journal of Animal Ecology
in 1932, and began studies leading to his important volume on animal populations,
Voles, Mice and Lemmings (Elton 1942).
The 1920s saw the beginning of the “Golden Age” of theoretical mathematical
population ecology. Raymond Pearl and L.J. Reed rediscovered the logistic equa-
tion describing population growth over time and advanced it as a “law of population
growth” (McIntosh 1985; Kingsland 1985). The logistic curve describes the growth
of a population over time as an S-shaped or sigmoid curve.1 The crux of the logistic
is that the rate of growth decreases as N increases and approaches K. The logistic
equation was expanded to the relations of two species, predation and competition,
independently by A.J. Lotka, a physicist, and Vito Volterra, a mathematician in the
1920s and later to n species. These ideas were pursued experimentally by a Russian
zoologist, G.F. Gause, working in the US for a period of years. These and similar
studies led to the formulation of “Gause’s Law” or the “competitive exclusion prin-
ciple” which was extensively pursued as ecological theory.
Mathematical population theory was expanded to n populations and was widely
criticized and praised as contributing to ecology, but in 1949 a volume on the
Principles of Animal Ecology (Allee et al. 1949) asserted that “theoretical population
1
It is commonly represented as dN/dt = rN (1-N/K) with r and K as constants, r being the maxi-
mum rate of dt K population increase in an unlimited environment, K the limiting population, N is
the number of individuals, t is time.
280
R. McIntosh
ecology has not advanced to a great degree in terms of its impact on ecological
thinking”. Nevertheless, mathematical ecology flourished as applied to competition
and predation. Here Robert May (1981) pointed out that the classical deterministic
logistic equation could under specific circumstances produce random looking dynam-
ics, not always a smooth sigmoid curve. In Australia, the ecologist Alexander J.
Nicholson and the physicist Victor A. Bailey believed that animal populations were
controlled by their density, or numbers, governed by competition for limited resources
(Nicholson and Bailey 1935). In the 1930s experimental studies were used for further
evidence for a balance or equilibrium of populations in nature. Disputes about popu-
lation control and theories thereof persisted in postwar ecology theory.
Australian ecology had begun in the 1920s largely as studies of plants and
animals with an economic basis. Ecology appeared in 1939 in a Conference of
the Australian New Zealand Association for the Advancement of Science. In 1951 the
Australian government agency, CSIRO, established a Section of Ecology, and an
Ecological Society of Society of Australia was formed in 1960.
Creating Larger Entities: Communities and Ecosystems
Study of aggregations of plants or animals, commonly called communities or asso-
ciations, was another aspect of protoecology and early ecology. Early British
marine biologists, called “dredgers”, studied marine bottom-dwelling organisms.
Edward Forbes (1844) recognized “zones” with species peculiar to them. Well
ahead of their time dredgers recorded location depths, species and numbers of indi-
viduals. A British biogeographer, H.C. Watson, advocated census of measured
areas (1 square mile) to determine the number of species present. His sage advice
was to increase the number of samples and decrease their extent. Fifty years later
Roscoe Pound and Frederic E. Clements did so, introducing the “quadrat” of one
square meter in larger numbers which was a major step in quantitative community
ecology (McIntosh 1985). In the smaller area it was feasible to count and measure
individuals. Plant ecologists had the advantage of stationary and more readily vis-
ible organisms. They came to recognize frequency as the number of samples in
which a species occurred, density as the number of individuals per area, cover as
the area of ground covered, and biomass (weight) as the measure of size. Studies of
the effect of number, size and shape of quadrats on these quantitative measures and
statistical analyses thereof occupied plant ecologists in subsequent decades (Greig-
Smith 1957). In 1949 G. Cottam and J.T. Curtis provided an alternative to the
quadrat by devising methods using distances from points in space.
Collection of data was followed by statistical analysis. In the US the efforts of
European ecologists, Paul Jaccard and O. Arrhenius, to examine the relation of
number of species and area were pursued by Henry Allen Gleason (1922). Gleason
(1920) had also examined the distribution of individual plants and used statistics to
test if they were dispersed at random but found they were most often distributed
contagiously in patches or non-randomly. The perception of communities as mosa-
ics or patchworks was advanced by William S. Cooper in the US and A.S. Watt in
281
20 The History of Early British and US-American Ecology to 1950
Britain (McIntosh 1985). Watt (1947) coined the apt term “gap-phase” for the
small-scale disturbances in communities.
Animal ecologists also pursued studies of animal communities, complicated by
the mobility and difficulties of sampling animals. Birge and Juday pursued their
studies of Midwestern lakes with sampling methods determining numbers of spe-
cies and individuals per area or volume and relating these to environmental mea-
sures. They provided what came to be an all too familiar lament of ecologists,
“The extension of our acquaintance with the lakes has been fatal to many interesting
and at one time promising theories” (Birge and Juday 1922). S.A. Forbes used
quantitative sampling methods and, in 1907, developed an early statistical index to
show co-occurrences of species among stands and conducted a survey of birds in a
cross section of the state of Illinois. Shelford studied insect communities on the
dunes of Lake Michigan and compiled a volume on Animal Communities in
Temperate America (Shelford 1913).
The British animal ecologist Charles Elton’s early work was largely on animal
communities and the interactions of populations in them. He considered the tradi-
tional analogy of the community as a clock but observed that the animal works
occasionally moved to another clock, calling into question the traditional balance-
of-nature concept and design metaphor of the clock (Elton 1930, pp. 16–17). He
later participated in a review of ecological surveys of animal communities (Elton
and Miller 1954) although much of his work was on populations.
In 1935 Arthur G. Tansley, in Britain, introduced a concept he termed “ecosys-
tem” in the context of a debate about F.E. Clements’ superorganism concept of com-
munity, then strongly advocated by South African ecologists and even Prime Minister
Jan C. Smuts who was influenced by Clements’ ideas. Tansley defined ecosystem as
the “whole system” including “the organismal complex” (biotic) and the “whole
complex of physical factors” (abiotic) called the environment. “Ecosystem” had
several antecedents, such as S.A. Forbes’ microcosm, and W.C. Allee’s geobio-
ecology in the English language literature, and others in European languages, sug-
gesting that the time was ripe for it. It fitted in with holistic traditions in ecology.
Tansley’s ecosystem incorporated a “hierarchy of systems of the most various kinds
and sizes” (Tansley 1935, p. 299). This generality persisted as a problem for ecosys-
tems ecology. “Ecosystem” came to be the major term encompassing the complex of
biotic community and physical environment and its later products, ecosystem ecol-
ogy and systems ecology, constituted for many a revolution in ecology.
Tansley’s term fell on well prepared ground, particularly in aquatic ecology. As
the president of the Ecological Society of America, W.P. Taylor (1935), had written,
“The emphasis placed by bio-ecology on organism and environment as a great uni-
tary problem is an inspiring one”. Among those inspired was a young limnologist,
Raymond Lindeman, who developed his studies of Minnesota lakes into a famous
article, “The Trophic-Dynamic Aspect of Ecology” (Lindeman 1942). Its departures
from the traditions of limnology were evident in that it was initially rejected by the
scientific journal Ecology on the basis of negative reviews by two distinguished
limnologists which were finally outweighed by the intervention of a third (Cook
1977). Lindeman used Tansley’s ecosystem concept, perhaps the first to do so, in
his “trophic-dynamic aspect” of ecology emphasizing the transfer of energy in the
282
R. McIntosh
ecosystem. Lindeman’s contributions were to emphasize and quantify trophic
function, establish a theoretical orientation in ecology and identify energy flow as a
fundamental process in long-term processes of community change (Cook 1977).
George E. Hutchinson, Lindeman’s mentor and supporter of his manuscript wrote,
in an addendum to the article, that Lindeman’s approach “may even give some hint
of an undiscovered type of mathematical treatment of biological communities”
(Lindeman 1942), although the paper’s critics were dubious about Lindeman’s data
and use of mathematics. Ecosystem ideas were the basis of Eugene Odum’s influen-
tial ecology textbook of 1953 which contributed to the introduction of the ecosystem
concept and its descendant, systems ecology, into ecology in the 1960s.
Another revolution in ecology was described by Barbour (1995) as appearing in
the 1950s. This revolution was against the tradition of the ecological community as
an organism or superorganism of integrated species developing to a stable climax
which was widely accepted in Anglo-American ecology (McIntosh 1998). The
revolution was instigated by belated recognition in the 1940s of three largely unrec-
ognized publications from 1917 to 1939 formulating the “individualistic concept”
of community. H.A. Gleason’s (1939) concept was based on his idea that each spe-
cies had individualistic properties, the environment varied in time and space, and
species aggregated into communities with a large stochastic component according
to vagaries of dispersal and the available environment. In 1947 Gleason’s long
ignored concept was resurrected by three prominent ecologists and in the 1950s
it was supported by extended studies of plants and animals by J.T. Curtis and
R.H. Whittaker, their students and associates. It became widely accepted by plant
and animal ecologists (McIntosh 1975, 1995). Even widespread acceptance of
Gleason’s concept and the ideas of continuum and gradient advanced by Curtis
and Whittaker did not entirely lay to rest the concept of equilibrium community and
even organism which persisted in theoretical ecology.
A Search for Laws and Principles
Coincident with concepts of population and community was a search for laws or
principles in early ecology. Both terms appeared with limited consensus on their
applicability in ecology. Victor Shelford (1913) introduced a “law of toleration”
which asserted that a species could grow in a range of environments with a mini-
mum, maximum and optimum represented by a single humped curve of distribu-
tion. Such curves were the substance of niche theory. Hopkins (1920) formulated
the “bioclimatic law” (Hopkins law) which recognized the familiar phenomena of
biological events moving north with the spring in the northern hemisphere. The law
stated that a biotic event lagged 4 days per degree of latitude in spring and early
summer, 5° of longitude eastward and 400 ft of altitude and gave rise to the aspect
of ecology, phenology, that followed such events. As ecology became more sophis-
ticated new laws and principles appeared. Preston (1948) described the distribution
of numbers of individuals of species distributed according to a “log-normal law”.
283
20 The History of Early British and US-American Ecology to 1950
Principles were more frequent and numerous. Gause (1936) offered an article on
“The Principles of Ecology”, Allee et al. (1949) indexed 25 principles and Odum’s
(1953) textbook in its 3rd edition published in 1971 recorded 30-odd principles.
Consensus on ecological laws and principles has been difficult to achieve and the
body of principles hoped for by Allee et al. (1949) did not readily emerge.
Nature as a Resource
Concern for nature and the conservation movement antedated ecology in Britain
and America. In Britain several public societies and government agencies were
formed in the 1860s to 1880s, and in 1894 a National Trust for the Preservation of
Places of Historic Interest and Natural Beauty was formed. In the US the 1870s saw
formation of a Fisheries Society and a US Fish Commission and the Audubon
Society. In 1872 Yellowstone National Park was formed and in 1892 the Sierra
Club was founded by the premier conservationist, John Muir, in time to defeat
attacks on the young national parks.
Many early ecologists were involved in the problems of land and wildlife manage-
ment and in conservation. A New Zealand ecologist, Leonard Cockayne (1918),
wrote that agriculture was applied plant and animal ecology. Aldo Leopold graduated
from forestry school, turned his attention to game management and ecology, and his
influential book, A Sand County Almanac, published posthumously in 1949, became
the classic statement of conservation in America. Paul Sears became a leading ecolo-
gist and exponent of conservation with his book, Deserts on the March (Sears 1935),
and his subsequent founding of a conservation program at Yale University.
In Britain Elton and Tansley were on a committee recommending a national
survey of soils and resources that was shelved due to wartime concerns, but during
the dark days of WW II Tansley and other ecologists served on a committee on
nature preserves. In 1949 the British Nature Conservancy was formed with Tansley
as its first chairman. By 1959 there were 84 nature reserves comprising 56.000 ha.
In the US Victor Shelford organized the Ecologists Union in 1946 that was trans-
formed into The Nature Conservancy which developed into the largest conservation
organization in the world.
Ecologists in Britain and America, well before the environmental crisis, recog-
nized its imminence and William Vogt (1948) published an early warning of the
crisis. Aldo Leopold (1949) extended ecology to ethics in his assertion, “That land
is a community is the basic concept of ecology, but that land is to be loved and
respected is an extension of ethics”.
Although “human ecology” was considered as early as 1913, the British
Ecological Society included it in its first meeting in 1914 and Clements (1935)
quoted H.G. Wells saying “Economics is a branch of ecology” and the Prime
Minister of South Africa saying “Ecology is for mankind”, widespread acceptance
of human ecology lagged. It was not until after WW II that ecology became widely
recognized as an integral component of human culture.
284
R. McIntosh
References
Adams CC (1913) Guide to the study of animal ecology. Macmillan, New York
Adams CC (1935) The Relation of general ecology to human ecology. Ecology 16:316–335
Allee WC, Emerson AE, Park O, Park T, Schmidt KP (1949) Principles of animal ecology.
Saunders, Philadelphia
Barbour M (1995) Ecological fragmentation in the fifties. In: Cronon W (ed) Uncommon ground:
toward inventing nature. Norton, New York, pp 75–90
Birge EA, Juday C (1922) The inland lakes of Wisconsin, the plankton 1. Its quantity and compo-
sition. Wis Geol Nat Hist Surv Bull 64:1–222
Clements FE (1905) Research methods in ecology. University Publishing Co., Lincoln
Clements FE (1916) Plant succession: an analysis of the development of vegetation. Carnegie
Institution of Washington Publ. 242, Washington DC, pp 1–512
Clements FE (1935) Experimental ecology in the public service. Ecology 16:342–363
Clements FE, Shelford VE (1939) Bio-ecology. Wiley, New York
Cockayne L (1918) The importance of ecology with regard to agriculture. N Z J Sci Tech
1:70–74
Cook RE (1977) Raymond Lindeman and the trophic-dynamic concept in ecology. Science
198:22–26
Cottam G, Curtis JT (1949) A method for making rapid surveys of woodlands by means of ran-
domly selected trees. Ecology 30:101–104
Cox DL (1979) Charles Elton and the emergence of modern ecology. Ph.D. dissertation,
Washington University, Washington, DC
Croker RA (2001) Stephen Forbes and the rise of American ecology. Smithsonian Institution
Press, Washington, DC
Egerton FN (1976) Ecological studies and observations before 1900. In: Taylor BJ, White TJ (eds)
Issues and ideas in America. University of Oklahoma Press, Norman, pp 311–351
Elton C (1927) Animal ecology. Sidgwick and Jackson, London
Elton C (1930) Animal ecology and evolution. Claredon Press, London
Elton C (1942) Voles, mice and lemmings: problems in population dynamics. Clarendon, Oxford
Elton CS, Miller RS (1954) The ecological survey of animal communities with a practical system
of classifying habitats by structural characters. J Ecol 42:460–496
Forbes E (1844) On the light thrown on geology by submarine researches. New Philos J Edinb
36:318–327
Forbes SA (1883) The food relations of the Carabidae and Coccindellidae. Bull Ill State Lab Nat
Hist 1:33–64
Forbes SA (1887) The lake as a microcosm. Bull Peoria Sci Assoc 111:77–87. (Reprinted Bull
Nat Hist Surv 15:537–550, Nov 1925)
Gause GF (1936) The principles of biocoenology. Q Rev Biol 11:320–336
Glacken CJ (1967) Traces on the Rhodian Shore. University of California Press, Berkeley
Gleason HA (1920) Some applications of the quadrat method. Bull Torrey Bot Club 47:21–33
Gleason HA (1922) On the relation of species and area. Ecology 3:158–162
Gleason HA (1939) The individualistic concept of the plant association. Am Midl Nat
21:92–110
Greig-Smith P (1957) Quantitative plant ecology. Butterworths Scientific, London
Hopkins AD (1920) The bioclimatic law. J Wash Acad Sci 10:34–40
Kingsland SE (1985) Modeling nature. Episodes in the history of population ecology. University
of Chicago Press, Chicago
Leopold AS (1949) Sand County Almanac. Oxford University Press, New York
Lindeman RL (1942) The trophic-dynamic aspect of ecology. Ecology 23:399–418
Mairet P (1957) Pioneer of sociology. The life and letters of Patrick Geddes. Humphries,
London
Malthus TR (1798) An essay on the principles of population. Johnson, London
285
20 The History of Early British and US-American Ecology to 1950
May RM (1981) The role of theory in ecology. Am Zool 21:903–910
McIntosh RP (1975) H.A. Gleason, “individualistic ecologist”, 1882–1975: his contributions to
ecological theory. Bull Torrey Bot Club 102:253–273
McIntosh RP (1985) The background of ecology: concept and theory. Cambridge University
Press, Cambridge
McIntosh RP (1995) H.A. Gleason’s ‘Individualistic Concept’ and theory of animal communities:
a continuing controversy. Biol Rev 70:317–357
McIntosh RP (1998) The myth of community as organism. Perspect Biol Med 41:427–438
Nicholson AJ and Bailey VA (1935) The balance of animal populations. Proceedings of the Zool
Soc Lond, 3: 551–598
Odum EP (1953) Fundamentals of ecology, 1st edn. Saunders, Philadelphia
Preston FW (1948) The commonness and rarity of species. Parts I and II. Ecology 43:185–218,
410–432
Sears PB (1935) Deserts on the March. University of Oklahoma, Norman
Shelford VE (1913) Animal communities in temperate America as illustrated in the Chicago
region. Bulletin of the Geographical Society of Chicago, Chicago
Tansley AG (ed) (1911) Types of British vegetation. Cambridge University Press, Cambridge
Tansley AG (1935) The use and abuse of vegetational concepts and terms. Ecology 16:284–307
Taylor WP (1935) Significance of the biological community in ecological studies. Q Rev Biol
10:291–307
Vogt W (1948) Road to survival. William Sloane Association, New York
Watt AS (1947) Pattern and process in the plant community. J Ecol 35:1–22
287
Chapter 21
The French Tradition in Ecology: 1820–1950
Patrick Matagne
A. Schwarz and K. Jax (eds.), Ecology Revisited: Reflecting on Concepts, Advancing Science,
DOI 10.1007/978-90-481-9744-6_21, © Springer Science+Business Media B.V. 2011
Conceptual history reveals that scientific ecology was built up around work
conducted during the nineteenth century by authors who often came from what
was, broadly speaking, a German cultural background. However, France also had a
presence in this field, in particular through authors who worked to develop the
study of botanical geography. Research in the history of ecology rarely explores
the literature of the learned societies in the French provinces. One reason for this is
that conventional historiography treats these groups and their work as social phe-
nomena that, however interesting to historians of French society, are assumed
a priori to be of little interest in terms of scientific content, except for local history
and archaeology, where their contributions are recognized.
In fact, a cross-analysis looking at the histories of ecology and these learned
societies together results in a contradiction of this conventional wisdom. The natu-
ralists of the French provinces, who are usually relegated to the margins of science,
can be seen instead to have helped to structure the scientific ecology.
Introduction
Since the 1960s, at a time when the general public was discovering the word “ecol-
ogy” in a context of growing awareness of the insidious effects of certain human
activities on the environment, historians of ecology in the United States and then in
Europe demonstrated that the founding concepts of scientific ecology had appeared
before the term was actually invented in 1866. They sketched out the conditions
underpinning the development of ecology as a scientific discipline based on work
conducted during the 19th century by authors who often came from what was,
broadly speaking, a German cultural background (Acot 1998). The institutional
period of ecology, characterized by congresses and university departments, began in
1900. During that period France stood out in particular due to the work of Augustin
P. Matagne (*)
Université de Poitiers, I.U.F.M., 40 avenue du Recteur Pineau, F-86000 Poitiers, France
e-mail: patrick.matagne@univ-poitiers.fr
288 P. Matagne
Pyramus de Candolle (1778–1841) – a Swiss man who worked in Montpellier – as
well as Gaston Bonnier (1853–1922) and Charles Flahault (1852–1935), who devel-
oped the study of ecological botanical geography (or phytogeography).
An entire domain of the scientific literature of the 19th century has, however,
remained unexplored: the impressive mass of publications of the learned societies.
This is due to several factors. The provincial learned societies had been gradually
relegated to the rank of associations of dilettantes who practiced science as amateurs.
This raises the issue of the relevance of the distinction between amateurs and
professionals. Unlike in the United States, where the main fields of science were
professionalized during the first half of the twentieth century, in France this matter
remained unsettled for much longer. The question remained in the nineteenth century
of what determined the boundary between the amateur naturalist and the professional:
was it the level and field of training, scientific style and method, institutional affiliation
(learned society, museum, library, laboratory, university, etc.) or the origin of any
remuneration? Was the professional distinguished from the amateur by their system of
theoretical reference, their working environment, their social standing, their way of
communicating or of disseminating science, or even by how close they were to Paris?
In practice, Alain Corbin observed that the capital “marked the successful cul-
mination of careers”. “At every level, Paris lived off the energy of the provinces, it
took in its men and tended to become for the provinces the paramount center of
culture, wealth and power, to the extent that the province evoked disgrace, remote-
ness from the center and a wasted life”.1 Another factor was a tendency in the
second half of the 19th century to see in the naturalist literature of the learned soci-
eties only unoriginal work, anecdotal or obsolete knowledge, the result of outdated
practices, leading to tiresome, pointless inventories. This was the view of the physi-
ologist Claude Bernard (1813–1878), who, on the occasion of the Universal
Exposition of 1867, which exposed the unsatisfactory state of affairs in France,
wanted to bring the natural sciences into the laboratory and the university. After the
debacle of the Franco-Prussian war of 1870, this desire to professionalize French
science, which was spurred by a spirit of revenge, led to hastening the disqualifica-
tion of amateurs. The learned societies were, as a consequence, relegated to the
margins of science (compare the situation in Spain at this time in Chap. 22).
As a victim of this dominant discourse, conventional historiography took an
interest in the contributions of these societies only in the fields of local history and
archaeology. It was not until the 1970s that their contributions to natural science
began to be evaluated without any a priori assumptions. It was in this spirit that a
parallel analysis was made between the history of the learned societies and the his-
tory of ecology (Laissus 1976, pp. 41–68; Dupuis 1979, pp. 69–106; Bange 1988,
1 “À tous les échelons, Paris se nourrit de la substance de la province, assimile ses hommes et tend
à devenir pour elle le centre primordial de culture, de richesse, de puissance, au point que la prov-
ince évoque la disgrâce, l’éloignement du centre et la moisissure de l’existence” (Alain Corbin
1992, pp. 793–794).
28921 The French Tradition in Ecology: 1820–1950
pp. 157–172). This revealed that the development of ecology did not merely take
place within conventional scientific institutions and that its actors were far from all
being professionals in the ordinary sense of the term. In addition, the day-to-day
presence of the naturalists in the local areas (natural environment, botanical garden,
laboratory, university, museum, library, biological station) and their membership in
a network of societies helped lay the basis for the organization of the first schools
of ecology in France, even before the well-known school of plant sociology in
Zürich-Montpellier. Examples of this include the École d’Auvergne, the École de
l’Ouest and the École méditerranéenne. These came together around the phytogeo-
graphic paradigm, but were divided by their different problematics.
The Naturalists in the Provincial Learned Societies
Out of the thousand provincial societies created between 1808 and 1914 (Fox, and-
Weisz 1980), about 350 conducted naturalist activity as part of their programme
(Fig. 21.1). The bulk of the societies’ publications consisted of works on flora and
fauna of local interest, as well as catalogues of plants and animals and monographs
and reports on outings. Their botanical work covered the entire territory of France
(Fig. 21.2). I conducted a survey on 28 societies (124 authors). This survey indicated
that botany was predominant (with 70%) over zoology (30%) (Fig. 21.3).
An analysis of this literature reveals certain aspects of the social dynamics of the
learned societies (Matagne 1997b, 1999a). Even if a work was signed by a single
author, many occasional contributors were cited or thanked collectively. For example,
the “Flore du Haut-Poitou” (1901) by Baptiste Souché (1846–1915), President of the
Deux-Sèvres botanical society, brought together contributions by botanists from the
society from 1889, as well as from their predecessors in the Deux-Sèvres statistics
society from 1870 onwards. James Lloyd (1810–1896), a botanist of English origin from
Nantes, cited 97 contributors in his “Flore de l’Ouest de la France” (1854). The pharma-
cist and director of the botanical gardens in Angers, Alexandre Boreau (1803–1875),
included 72 contributors in his “Flore du Centre de la France” (1857).
Information and samples were exchanged in the field, and others were sent by
post. A variety of species were circulated for the purpose of verification, or as
exchanges or gifts. Large herbaria and collections of insects, rocks, stones and
shells were offered, sold or bequeathed in whole or in part.
The societies’ meetings gave rise to reports listing the species discovered. The societ-
ies encouraged and handled exchanges and published lists of available samples, opening
what amounted to trading centers. They even founded groups devoted exclusively to this
activity, such as the Pyrenees association for the exchange of plants, founded in 1890.
Consider the case of the Deux-Sèvres botanical society. Founded in November
1888, by the next year it already had 153 members, and over 350 members by the
turn of the century. It became a regional botanical society, with 632 members on
the eve of the First World War. It organized 1,254 outings between 1889 and 1914.
290 P. Matagne
Entomology
*some articles are written by several authors
SocietiesAuthorsBotany ZoologyPalaeontology
Pluridisciplinary:13
839 24 14 10
53* 30 13 00
Natural sciences:
Agriculture: 3
2
2
28 124 70 32 11
6400 0
54001
21 850 0
Geography:
Botany:
Total:
Fig. 21.2 Number of authors of floristic books and catalogues (1800–1914)
Fig. 21.1 Geographical distribution of scientific societies which practiced natural history
29121 The French Tradition in Ecology: 1820–1950
In a single outing, 200 species were collected. In his field notes, Souché noted that
on 17 May 1886 he had collected 160 samples of the same species. One month
later, he filled his famous green botanic sample box (Fig. 21.4). In September of
that same year, he prepared 52 samples of each of 12 species for the La Rochelle
natural sciences society. The naturalists, whose curiosity was pushing them towards
less accessible areas (the Alps, the Pyrenees), viewed their mission as collecting
large quantities of rare species (Matagne 1988, 1997a) – an attitude that would be
considered shocking in the twenty-first century! But at that time naturalists saw
themselves as hunters of plants and animals.
The naturalists were long regarded as representatives of an outdated science, due
to their passion for collection, classification and inventories. Their reports on their
festive, sportive outings, events marked by their social character, replete with pic-
nics, toasts, debates and anecdotal exchanges, could lead to confusion between the
activities of these learned societies and those of other societies, such as singing or
athletic groups. But promoting this social side had strategic value, as it attracted
Fig. 21.3 Number of naturalist’s publications of 28 learned societies in the “province”
292 P. Matagne
Fig. 21.4 (a–e): Basic tools of the botanising naturalist. (a) Herborization box, (b) and (c) tools
for digging and breaking, (d) “échenilloir” et “sécateur”, (e) knapsack (Figures taken from
Bernard Verlot 1879. Le Guide du botaniste herborisant. - Paris: J.-B. Baillière et fils; Guillaume
Capus 1883. Guide du naturaliste préparateur et du voyageur scientifique, ou Instructions pour la
recherche, la préparation, le transport et la conservation des animaux, végétaux, minéraux, fossiles
et organismes vivants. 2e édition, entièrement refondue par A.-T. de Rochebrune, avec une intro-
duction par E. Perrier. Paris: J.-B. Baillière et fils)
people whose dues replenished the often meager funds of the groups. The newsletters
also of course published work that reflected the interest of the learned societies in
botanical geography.
The Practice of Botanical Geography
Nowadays, botanical geography is the study of the factors that influence the
distribution of plants, the most important of which are geology, the climate, and the
mechanism for the dispersion of the reproductive organs of the plant itself.
Phytogeography thus includes the causal study of the distribution of species, the
cartography of information showing the natural distribution of plants, and an index
of the plant species and associations of a given region.
Baron Alexander von Humboldt (1769–1859), a German naturalist and explorer,
is considered one of the founders of phytogeography. In 1805, he initiated an
approach to landscapes that aimed at establishing phytogeographic classifications
based on physiognomic analogies (Humboldt 1805). For example, the beech groves
on the massif of the Grande Chartreuse (the Dauphiné area of the Alps) and the
high-altitude beech groves in New Zealand and those in Croatia (below 1,700 m)
29321 The French Tradition in Ecology: 1820–1950
Fig. 21.4 (continued)
have a similar, characteristic physiognomy determined by the beech tree, even
though the floristic composition of the plant community dominated by the tree varies
with latitude and altitude.
The Swiss botanist Augustin Pyramus de Candolle originated the floristic tradi-
tion in phytogeography. The identification of plant communities is determined
based on the most comprehensive comparison of inventories possible, which makes
it possible to identify the species typical of associations that are always at least
statistically measurable present in a specific environment. There, they could have
only a minor impact on the physiognomy of the landscape. This is true for example
of Scilla lilio-hyacinthus (Squill lily-hyacinth), which is characteristic of the asso-
ciation of the submountain beech groves with firs in Auvergne, which are found
near Lake Pavin.
294 P. Matagne
Fig. 21.4 (continued)
29521 The French Tradition in Ecology: 1820–1950
The two traditions were developed by authors who attempted to develop a
taxonomy that has the association as its basic unit (Matagne 1998). These traditions
persisted through the century and formed part of the foundation of ecological
botanical geography, the first treatise of which was published by the Danish bota-
nist Eugenius Warming (1841–1924) in Danish (1895) and German (1896), and
then in English (1909) (see for more details see Chap. 23). This was the era of the
first generation of European and North American ecologists.
As early as 1810, the botanists from the Orléans Society for the Physical and
Medical Sciences took an interest in the climatic and geological causes of the variation
in flora between the Paris and Orléans regions. References to phytogeography became
generalized in the 1820s, as was shown by the survey mentioned earlier which com-
prised 18 French départements (25% of the societies surveyed; Fig. 21.1).2
Phytogeographers studied the relations of plants with the type of terrain, tempera-
ture, altitude, and exposure in order to determine the laws governing their distribution.
Sometimes the impact of human activity was considered. Starting in the 1850s, a period
that saw the appearance of “Géographie botanique raisonnée” (de Candolle 1855) by
Alphonse de Candolle (1806–1893) and “Etudes sur la géographie botanique de
l’Europe” (Lecoq 1854) by Henri Lecoq (1802–1871), anyone who did not deal with
these issues was marginalized by the Aveyron society of letters, sciences and the arts.
The survey showed that the new concepts in botanical geography, whether origi-
nating with Candolle or Humboldt, made sense out of discoveries that had thereto-
fore caused confusion: the abnormal presence of Pteris aquilina (bracken fern) on
limestone was cleared up if one invoked the nature of the bedrock, exposure and
certain local atmospheric factors. Plants became environmental indicators: compiling
a list of species was sufficient for the recognition of a limey soil. The phytogeo-
graphic paradigm served as the key to floristic transitions, the preferences of a
given species, and the causal study of areas of distribution.
A.P. de Candolle drew the attention of naturalists and agronomists to potential
applications (1809, pp. 335–373). Agricultural societies opened their newsletters to
naturalists and chemists. Behind questions of yields, selections and attempts at
acclimatization lay ecological issues concerning the relations between the plants
and the soil (Dagognet 1973, pp. 51–52). References to Humboldt and to de
Candolle, father or son, were rarely explicit. These could be identified rather by the
use of various concepts, such as the “station”, which was employed in different
ways by Humboldt and by A.P. de Candolle (Drouin 1991, pp. 74f).
The botanists also undertook to identify and classify associations.3 This research
on associations changed profoundly botanical practice. In the field, the botanists,
having identified species that were always associated with the same environment,
attempted to determine what accompanied them: the concept of association helped
to identify regularities in nature, to make and then verify predictions, and to identify
2 Ain, Aisne, Ardennes, Aude, Aveyron, Cher, Doubs, Haute-Garonne, Gironde, Indre, Indre-et-Loire,
Loire-Inférieure (actuelle Loire-Atlantique), Loiret, Maine-et-Loire, Saône-et-Loire, Deux
Sèvres, Vendée, Vienne.
3 The notion “association” was formulated for the first time by Humboldt in 1805.
296 P. Matagne
anomalies and search for their causes. For example in 1912, Emile Château
(1866–1952), a teacher and then primary school head, published an innovative
work that has nevertheless remained relatively obscure, entitled “Les associations
végétales”, which was a precursor of plant sociology developped by the Zürich-
Montpellier school in the 1920s (Château 1912, pp. 175–192). Others referred to
“the concept of plant formation” (1838), formulated by the Göttingen botanist
August Grisebach (1818–1879). In the tradition of Humboldt, a plant formation
was understood as a group of plants with a definite physiognomy. Finally, some
explored the physiognomic-floristic path laid out in 1863 by Anton Kerner von
Marilaun (1831–1898). The Austrian botanist focused on floristic composition and
on the physiognomy of plant groups, and began to free them from the environment,
defining them in their own right, as in the case for species (von Marilaun Anton
1863). The consensus established around the phytogeographic paradigm showed
the capacity of amateur naturalists to take on board rapidly the latest scientific
contributions, originally humboldtian and candollian, incorporate them into phyto-
geographical practice and to adapt them to the local problems. Their attempts at a
synthesis between the physiognomic and floristic traditions, and their search for a
definition of an association that was considered sometimes as determined by the
environment and at other times as a separate entity, all provided material for plant
ecology during the 1920s and 1930s. This also anticipated the rupture that the
Zürich-Montpellier school would make with phytogeography. All in all, the learned
societies harbored pioneers in the ecological botanical geography of France. They
were active in a number of fields where they had gained scientific credibility.
The Fields of Ecology
The natural environment is the terrain for botany par excellence. There are, how-
ever, other areas where naturalists exercise their skills and talents, geographical
ones of course, but also institutional and social ones, such as gardens, museums,
and biological stations.
During the nineteenth century, almost every large city created its own garden,
under the pressure of the universities and learned societies, who wanted to be in
charge of their administration.
Traditionally, the arrangement of plants was systematic (based on a classifica-
tion system), so that one speaks of a “school of botany”. The systems of Joseph
Pitton de Tournefort (1656–1708), Carolus Linnaeus (1707–1778) and the methods
of Antoine-Laurent de Jussieu (1748–1836) and A.P. de Candolle (1818–1821) suc-
ceeded one another or coexisted along corridors of labelled samples.4 The arrange-
ment of plant beds thus presented a summary of the geographic distribution of
4Joseph Pitton de Tournefort, Éléments de botanique ou Méthode pour connaître les plantes
(1694); Carolus Linnaeus, Genera Plantarum (1737); Antoine-Laurent de Jussieu, Genera
Plantarum (1789) and A.P. de Candolle, Reyni vegetalis Systema naturale (1818–1821).
29721 The French Tradition in Ecology: 1820–1950
plants. The societies promoted the use of these plant beds, as in Bordeaux (1847),
and later in Béziers (1886) and Niort (1890). At the end of the century, some people
wanted to create ecological gardens where the plants were arranged in their cus-
tomary situation, in varying exposures Tarn (1882–1883). These gardens respected
botanical affinities and highlighted the plant characteristics, importing the soil from
each different region of origin Clermont-Ferrand (1893).
Alpine gardens, like the one started at the Pic du Midi observatory in 1878, were
models. Edouard André, a landscape architect influenced by von Humboldt, argued
that it was possible to reconstitute mountain associations in the plain. The French,
following on the heels of the English, Swiss and Germans, attempted more or less
successfully to reconstitute rock gardens (André 1894, p. 1228; Matagne 1992,
1999b, pp. 307–315). The various conceptions held by local scholars were reflected
in the school garden (systematic), the ecological garden (relationship with the envi-
ronment) and the geographic garden (by region of origin).
The passion for museums (traditional, revolutionary or commercial), often in asso-
ciation with libraries, reflected the rise of a new way of looking at the natural heritage
bequeathed by the French Revolution. Here too the local societies were in demand,
particularly since the means and skills needed to identify, classify and preserve the
diverse collection of innumerable kinds of objects were lacking. In terms of natural
history, the goal was comprehensiveness at the local level. Nîmes (1822), Nice (1828),
Rouen (1834), Poitiers (1836), Le Havre (1845), Bayonne (1856), and Toulouse (1865),
etc., all had their museum of natural history, created or supported by local societies.
The gardens, museums and libraries came to reflect the learned character of the
societies that took responsibility for them. The last third of the nineteenth century
then witnessed the boom in agronomic and botanical stations. Learned societies
seized the opportunity to modernize by playing a role in these stations, with their
modern laboratories (Matagne 1996).
Agronomic stations were established in France relatively late compared with
Germany (Boulaine 1992). They had experimental fields and cattle pens as well as
public laboratories. The technical methods used in chemistry, physics and physiology
were harnessed to help agriculture. Farming societies linked to the stations promoted
laboratory analysis and took part in developing agronomic maps. They opened their
doors to naturalists by creating natural history sections.
Maritime stations were set up during this same period and opened their doors
to amateurs. By the time of the First World War, the French coast had a dozen of
them (Fischer 1997). One noteworthy case was the station at Arcachon, which was
established in 1867 and for 30 years was owned and administered by the Arcachon
scientific society. Innovative research programmes were launched in marine
biology, oceanography, physiology and electrophysiology, and in the field of the
ecology of the dune environment.
Also worthy of note is the station at Mauroc, established on 30 May 1912 by the
University of Poitiers. For the next year the regional botanical society headed by
B. Souché was extremely active meeting demand for the organization of outings
and the creation of a botanical garden, a library and herbaria. In exchange, amateurs
were invited into the laboratories.
298 P. Matagne
Contrary to the general view, the learned societies thus had the desire and the
capacity to take on board new scientific concepts, to apply them at various levels, and
to familiar their members with laboratory practices. They were thus not pushed to the
periphery of scientific inquiry. They even played an innovative role by taking charge
of the promotion of ecological gardens and creating the first schools of ecology.
The First Schools of Ecology
The “École du Centre”, the “École de l’Ouest” and the “École Méditerranéenne”
reflected a diversity of approaches related to particular phytogeographic, sociologi-
cal and local institutional features.
According to the phytosociologist Jules Pavillard (1868–1961), Auvergne was the
“classic area of French phytogeography” (Pavillard 1926, p. 2). He is referring to the
“Etude sur la géographie botanique de l’Europe et en particulier sur la végétation du
Plateau central de la France” (1854) by Henri Lecoq, professor of natural history and
director of the Clermont-Ferrand botanical garden. A number of well-known figures
had spent time in Auvergne before Lecoq, including the botanists René-Louiche
Desfontaines (1750–1833) and Adrien de Jussieu (1797–1853), the Pyrenees specialist
Louis Ramond de Carbonnières (1755–1827), A.P. de Candolle, the explorer of India
and the Himalayas Victor Jacquemont (1801–1832), the explorer of the Balearic Islands
Jacques Cambessedès (1799–1863), the travelling botanist Auguste Prouvensal de
Saint-Hilaire (1779–1853), the founder of modern geology Charles Lyell (1797–1875),
the Montpellier professor Charles Flahault, the authors of Flores françaises Gaston
Bonnier and Father Hippolyte Coste (1858–1924), and many others. A stopover in
Auvergne seemed a must for any naturalist travelling around France or the world. Many
amateurs published in the “Annales scientifiques”, which had been founded by Lecoq
and placed in the hands of the Académie at Clermont-Ferrand, which did not remain
very active. Lecoq had difficulty establishing a following in his region before the cre-
ation of the natural history society in Auvergne (1894), which made use of his work.
From the 1820s, Lecoq, a follower of Humboldt, called on botanists to focus their
work on plant associations, “these natural groups that form the subject matter of
botanical geography”.5 He developed an original geological and paleontological
approach. In a catalogue produced with his student Martial Lamotte (who died in
1883), he set out methodological guidelines for the readers of the “Annales scienti-
fiques”: “the geological study of such a varied soil made it much easier for us to
resolve the previously unanswered question of the terrain’s influence on plant
stations”.6 He recommended breaking down the Massif Central into more limited areas
based mainly on geology and secondarily on topography and plant physiology.
5 “ces groupes naturels, dont l’étude constitue la géographie botanique” (Lecoq 1854, p. X).
6 “l’étude géologique d’un sol aussi varié nous a donné de grandes facilités pour résoudre la ques-
tion jusqu’ici indécise de l’influence des terrains sur les stations des plantes” (Lecoq and Lamotte
1847, p. IX).
29921 The French Tradition in Ecology: 1820–1950
Inspired by Humboldt’s approach with regard to “the sociability of plants and
their associations between similar individuals and between different species”,
Lecoq drew up a scale of dominance to evaluate the presence of one or another
species, with a view that “the gathering of all the species of a single station consti-
tutes a plant association”.7 This scale was used to distinguish dominant, essential,
secondary and accidental species. Another student, S.E. Lassimonne, then drew on
this for his study “principes de topographie botanique” (1892), which distinguished
between isolated and associated species. Lecoq also developed an algebraic equa-
tion that described the relationship between the physico-chemical characteristics
(atmospheric, edaphic) of a station. This attempt at establishing quantitative or
mathematical relationships was a first in ecology.
A botanist from Aveyron, Joseph Revol, drew on Lecoq for his study of the flora
of Southwestern France. So too did Charles Bruyant for his research into the evolu-
tion of plant groups in the lakes and peat bogs of the Mont-Dore area. Bruyant, a
professor at the Clermont-Ferrand medical school, attempted to apply this method-
ology to zoogeography. Two works that dominated regional botany and phytogeog-
raphy for a half-century were at the source of another school in the West of France
“Flore de la Loire-Inférieure” (1844) and “Flore de l’Ouest” (1854), by James
Lloyd. He had lived in France since the age of six, finished high school at Lorient,
and then moved to Nantes and finally, from 1840, to Thouaré-sur-Loire (Loire-
Inférieure, now Loire Atlantique).
In 1844, Lloyd argued that it was necessary for botanists to go beyond their
administrative boundaries, and he urged them to study successions, striking exam-
ples of which were abundant along the Atlantic coastline. This abundance was due
to a highly selective climate in the sea spray area. Lloyd noted that the strong winds
and brisk, salty air determined the small-scale species, with fleshy leaves and thick,
almost ligneous roots that dug deep into the soil in their search for fresh water.
Lloyd’s phytogeographic programme consisted of comparing the coastline flora
with that of the interior so as to identify transitions:
[...] if we start from the southern extremity of the coast, we find the salty marshes of
Bourgneuf (area of Retz, Brittany marshland), the leading station for plants specific to salty
silt[...]”; “[...]it is always useful to examine the points where the salt marshes arise, mixing
with cropland, sand, marshy pastureland and fresh water streams. At Collet begins the
maritime sand and the plants specific to it [...]”; “[...] before leaving the maritime area, [...]
I would recommend that research botanists look at this part of the coast, which forms the
transition between the maritime flora and the flora of the interior.8
7 Lecoq 1854, p. 134.
8 “[...] si nous partons de l’extrémité méridionale de la côte, nous trouvons les marais salants de
Bourgneuf (pays de Retz, Marais Breton), première station des plantes propres aux vases
salées[...]”; “[...] il est toujours utile d’examiner les points où les marais salants prennent
naissance, en se confondant avec les terres cultivées, les sables, les pâtures marécageuses et les
ruisseaux d’eau douce. Au Collet commencent les sables maritimes et les plantes qui lui sont
particulières[...]”, “[...]avant de quitter la région maritime, [...] je recommanderai aux recherches
des botanistes cette partie de la côte qui forme la transition de la flore maritime à celle de
l’intérieur” (Lloyd 1844, pp. 14–24).
300 P. Matagne
Ten years later, Lloyd’s Flore de l’Ouest covered eight départements. The boundaries
were determined by the nature of the soil and climatic, physiognomic and physio-
logic factors. Lloyd had established his presence in the area and had numerous
pupils in the learned societies of La Rochelle, Morbihan, Finistère, Deux-Sèvres
and Loire-Inférieure, where the academic society officially introduced his research
programme.
The son of a grocer, Emile Gadeceau (1845–1928), reviewed ecology’s treat-
ment of the causes of floristic and physiognomic successions in light of the con-
cepts of American ecology. As a member and then president of the Loire-Inférieure
academic society, he published studies influenced by Lloyd and Flahault. His most
innovative work was entitled “Le Lac de Grand-Lieu. Monographie phytogéographique”
(Gadeceau, Èmile 1909). The lake, located some 15 km southwest of Nantes,
has been a nature reserve since 1980. There were three parts to Gadeceau’s work:
lake geography, aquatic plants, and biological ecology. He made use of the concepts
of Warming and the Strasbourg botanist Wilhelm Schimper (1856–1901), who laid
the physiological basis for plant ecology that Warming lacked (Schimper and
Andreas Franz 1898).
The third part of Gadeceau’s “Essai” introduced American notions of ecology for
the first time. Starting from Lloyd’s spatial view of succession, Gadeceau adopted the
approach of the official botanist of the state of Minnesota, Conway MacMillan (1867–
1929), who in 1897 studied plant distribution on the banks of a lake from a dynamic
viewpoint. The point was to follow successions not only over space but also over time.
Gadeceau also cited Henry Chandler Cowles (1869–1939), from the University of
Chicago, who published a thesis on Lake Michigan (Cowles, 1899). The two authors
were among the pioneers of the Chicago school of ecology (see Chap. 20).
Gadeceau read the Botanical Gazette and the Monde des Plantes published in
Mans in the Sarthe region – a mine of bibliographical references – and was familiar
with the latest thinking and applied it to the case of the Grand-Lieu Lake. His epis-
temological position was thus remarkable: an amateur botanist from a provincial
learned society had introduced North American ecology into France before the First
World War. The École de l’Ouest identified with Lloyd had of course created
theoretical conditions that were propitious for integrating this new ecology, but this
would still require moving from the conventional photographic vision of European
phytogeography to America’s cinematographic vision.
The first works on botanical geography in the Mediterranean region were carried
out by Father Giraud Soulavie (1752–1813) (Soulavie 1780–1784), who evoked the
value of territorial divisions based on climatic factors. Father Pourret (1754–1818)
contributed material on the Eastern Pyrenees, and Marquis Gaston de Saporta
(1823–1895) drew attention to the importance of the history of flora for an explana-
tion of its current state.
The “Essai d’une géographie botanique du Tarn” published in 1862 was a pre-
cursor, even though it failed to gain a following (Gazel Larambergues 1862) as it
set out an innovative programme that proposed to divide the département on the
basis of topography and the nature of the soil rather than on the traditional basis of
administrative limits (arrondissement, canton, commune).
30121 The French Tradition in Ecology: 1820–1950
In the neighboring département of Tarn-et-Garonne, Dominique Clos (1821–1908)
laid the basis for the Mediterranean school of ecological phytogeography during a
local congress in 1863. His programme was published in 1864 in Toulouse and in
1870 in Carcassonne (Clos 1864, 1870). Clos proposed seeking the limits of
Mediterranean vegetation towards the west of Carcassonne and the south of Tarn
by tracing the disappearance of certain species and sketching out their areas. The
Carcassonne society of arts and sciences adopted this programme and identified the
natural divisions within the département, which demarcated the Mediterranean
phytogeographic region. Two contemporaries of Clos, the Hérault botanists Henri
Loret and Auguste Barrandon, made use of this programme in their Flore de
Montpellier (Loret and Barrandon 1876).
Clos, Loret and Barrandon thus were initiators of a Mediterranean school that
adopted and adapted the phytogeographic paradigm based on Candolle. Their start-
ing point was the principle that a département-based breakdown offered an easy and
relevant framework of grids that botanists could then use to seek the causes of plant
distribution and identify associations. They expected that when every département
had its phytogeographic map, the puzzle would be complete, including for the
country as a whole. Hector Leveillé (1863–1918), the head of “Le Monde des
Plantes”, supported Loret’s quest.
The Béziers Society for the study of the natural sciences joined the programme,
followed by the Aude Society for scientific studies. Publications came out up to
1909 that referred directly to Loret and Barrandon “Catalogue des plantes vasculaires
du département de l’Ardèche” (Baichère 1891; Gautier 1898; Albert and Jahandiez
1908). Finally, a dozen authors out of the 50 who mattered in the region
(Fig. 21.3) trained the botanists of a number of learned societies.
Nevertheless, Professor Charles Flahault took a dim view of this ecological
work, which broke with the traditional practices of non-causal descriptive
phytogeography (chorology). Even though he acknowledged that Mediterranean
phytogeography was on the right path, he deplored the départemental confines
that it set itself. He believed, for instance, that work on the flora of the lime-
stone plateaus of the Causses region would be of greater interest than that in the
départements of Aveyron and Lozère. He condemned those who studied “plants
that have nothing in common other than being under the jurisdiction of the
same civil servant”.9
Flahault had in fact chosen the Mediterranean region as the grounds for an ambi-
tious study. He wanted to make use of the local societies to spread his programme,
which consisted of determining the limits of the natural areas of plants, without
worrying about administrative boundaries. In other words, he wanted the learned
societies to work on behalf of his programme, with a view to developing a carto-
graphic summary at a regional and then national level.
9 “des plantes qui n’ont en commun que d’être sous l’administration d’un même fonctionnaire”
(Charles Flahault 1901, pp. 3–4).
302 P. Matagne
Flahault’s renown, his personality and his campaigning style had an influence
well beyond the regional level. He took part in the debates and controversies raging
among phytogeographers at the turn of the 19th and 20th centuries. It was urgent at
that time to develop a consensus on phytogeographic nomenclature and on the defi-
nition of an association, in order to avoid a “Babelization” of the discipline
(Bonnier 1900). Ultimately, it was the floristic line around which the Zürich-
Montpellier school had organized that won the day in continental Europe. It defined
an association in a way that ruptured with the approach of the phytogeographers:
the association was to be identified by its floristic composition, and not by environ-
mental factors.
Flahault made an appearance in 1888 at a conference in the Narbonne area of the
French botanical society, where he met with people capable of spreading his ideas.
In 1889, he helped set up the Aude scientific society, whose botanists officially
declared that they were abandoning administrative limits. Twenty-five authors (out
of 250 members) in the society published work along these lines. The already-
mentioned rather modest society in Carcassonne went over to Flahault’s approach
in 1890–1892. The Languedoc geographical society, which was linked to the
botanical institute founded by Flahault in Montpellier, made a powerful contribu-
tion to spreading his ideas.
Various journals ensured the promotion of Flahault’s approach beyond the
region, including the Revue scientifique du bourbonnais, Monde des Plantes, and
the Bulletin de la société botanique de France. Distant followers of Flahault were
active, such as Eugène Simon (1871–1967) in the Deux-Sèvres area (Matagne
1990). Finally, Flahault wrote an introduction to the scholarly but popular “Flore de
France” by Father Coste (1901–1906), in which he promoted his programme
(Flahault 1901). In the absence of centralized, standardized data, it is difficult to
arrive at an assessment of the work of the school that preceded Flahault. A tenth to
an eighth of French territory could have been covered. Flahault considered the
results unusable. A review of publications from the period 1920–1930 indicates the
need for another analysis.
The Flahault school, symbolically founded in 1888, brought together some 30
authors. In 1909, they had a cartography of a tenth of the country. Despite this,
Flahault gave up due to a lack of support and of funds. He was unable sufficiently
to mobilize the local societies, who resisted a form of instrumentalization and pre-
ferred to stick to tried and proven methods of exploration. Moreover, phytosociol-
ogy came onto the plant ecology landscape scene at the start of the 20th century and
resolved the problems in terminology and nomenclature in which phytogeography
had become bogged down.
Other schools also undoubtedly existed, in particular around Toulouse and in
the northeast of France.10 The three that have been presented here illustrate the
epistemological work of the Catalan ecologist Ramón Margalef. He notes that
10 In Toulouse this was Lavialle before 1914, then Henri Gaussen; in the north-east, Alexandre Godron
(1807–1880) and Paul Fliche (1836–1908) contributed to phytogeography, zoogeography and forestry.
30321 The French Tradition in Ecology: 1820–1950
ecology is marked by a sort of Genius loci, which would have the dynamics of
schools linked to local landscapes: hence the phytosociology school, which devel-
ops concepts and methods adapted to the mosaic of Alpine and Mediterranean
vegetation, and North American ecology, which is absorbed in the gradual tran-
sitions of large spaces, plant successions and pioneering species (Margalef 1968,
p. 26). Lloyd and Gadeceau encountered this type of environment on a smaller scale
along the French Atlantic coastline. An analysis of the environment in Auvergne
shows the importance of geological factors. In the Mediterranean area, the eco-
logical problem encountered the political problem of the administrative limits set
by the French Revolution, which determined those of the small provincial “moth-
erland” to which the local scholars were so attached.
Conclusion: The Unique Features of French Ecology
The destiny of the first schools of ecology was linked to the fate of the learned societies
that organized and supported them. Many society members perished in the First World
War. Greater labor mobility also hindered the development of local roots. In addition,
devaluation ruined many learned societies, which often depended on investments.
Those that re-formed during the 1920s did so on a different basis. The first schools of
ecology thus disappeared with the learned societies before the First World War.
Nevertheless, it was not uncommon to see references to their authors in the
1920s. Pierre Allorge (1891–1944), professor of cryptogamie at the Museum of
Paris, used the work of Lecoq in his phytosociology study “Associations végétales
du Vexin français” (1922), in a chapter reviewing the accomplishments of ecology.
There are also specific references to Lecoq in the work of Aimé Luquet, “Essai sur
la géographie botanique de l’Auvergne”, subtitled “Associations végétales du
Massif des Monts Dores”, which also cites the work of Lamotte and Bruyant, who
followed in Lecoq’s footsteps. Nevertheless, in this “Essai”, dedicated to Braun-
Blanquet and to Pavillard, it is the contributions of the phytosociological school
that Luquet belonged to that are given greatest emphasis. Even so, there are still
references to 19th century authors from Auvergne in “Recherches sur la géographie
botanique du Massif Central”, also by Luquet, in 1937.
Lloyd and Gadeceau are referenced in botanical and phytogeographical studies.
Nevertheless, the introduction of concepts from American ecology, discovered
before 1914 by Gadeceau, continued its course with other authors in the 1920s and
1930s, including Allorge, Emmanuel de Martonne (1873–1955) and Henri Gaussen
(Acot and Drouin 1997).
Plant ecology in the 1920–1950 period came to be marked in France by the
phytosociology propounded by Braun-Blanquet. It is also characterized by bio-
cenotics and by the unique approach to human ecology that grew out of the geog-
raphy of Vidal de la Blache (1845–1918).
Due to the domination of phytosociology, for the participants in the first inter-
national colloquium of the “Centre nationale de la recherche scientifique” (CNRS)
304 P. Matagne
on ecology, held in Paris from 20 to 25 February 1950, it was important to draw
attention to the study of animal groupings (animal synecology), which was lagging
behind. Even though phytosociology was thus officially not on the agenda, it was
omnipresent. On the other hand, the theory of ecosystems, then all the rage in the US
(Odum and Odum 1953), was missing. According to Acot Pascal (1994), a historian
of ecology, this reflected a unique feature of French ecology in the 1920–1950 period.
Another unique feature of the French tradition in ecology had its roots in the
19th century. It was born and developed through the work of countless authors
whose names conceptual history has not retained. They contributed to developing
a French approach to ecology based on the phytogeographic paradigm organized
around schools distinguished by their local dynamics.
At the beginning of the twentieth century, the institutionalization of ecology, the
First World War, the rise of phytosociology, and the sudden emergence of American
ecology led to the marginalization of these local schools of ecology. What remained
of them after the War was a dispersed heritage and a few individual authors who
were decontextualized by those who later cited their works.
Fig. 21.5 The map displays all the locations of early French ecology activities mentioned in the
article (arranged by C. Haak)
30521 The French Tradition in Ecology: 1820–1950
References
Acot P, Drouin J-M (1997) L’introduction en France des idées de l’écologie scientifique américaine
dans l’entre-deux guerres. Revue d’histoire des sciences 50:461–479
Acot P (1994) Le colloque international du CNRS sur l’écologie (Paris, 20–25 février 1950). In:
Debru C et al (eds) Les sciences biologique et médicales en France, 1920–1950. CNRS édi-
tions, Paris, pp 233–240
Acot P (ed) (1998) The European origins of scientific ecology. Gordon and Breach Publishers,
Amsterdam
Albert A, Jahandiez E (1908) Catalogue des plantes vasculaires qui croissent naturellement dans
le département du Var. Paul Kliensieck, Paris
André E (1894) Les Fleurs de pleine terre comprenant la description et la culture des fleurs annuelles,
bisannuelles, vivaces et bulbeuses de pleine terre. Vilmorin-Andrieux et Cie, Paris, p 1228
Baichère A (1891) Contributions à la flore du bassin de l’Aude et des Corbières. Bulletin de la
société d’études scientifiques de l’Aude, p 73
Bange C (1988) La contribution des ecclésiastiques au développement de la botanique dans la
région lyonnaise au 19e siècle. 112e Congrès national des sociétés savantes, Lyon, 1987.
Section histoire moderne et contemporaine, Paris, pp 157–172
Bonnier G (1900) Projet de nomenclature phytogéographique. Actes du 1e Congrès International
de Botanique tenu à Paris à l’occasion de l’Exposition Universelle de 1900. E. Perrot,
Lons-Le-Saunier, pp 427–449
Boulaine J (1992) Histoire de l’agronomie en France. Technique et Documentation, London,
New York, Paris
Boreau A (1857) Flore du Centre de la France. Librairie Encyclopédique de Roret, Paris
de Candolle A-P (1809) Géographie agricole et botanique. In: Nouveau cours complet d’agriculture
ou Dictionnaire raisonné et universel d’agriculture, vol 6. Déterville, Paris, pp 335–373
de Candolle A-P (1818) Reyni vegetalis Systema naturale. Treuttel & Würz, Paris
de Candolle A (1855) Géographie botanique raisonnée. Libraire de Victor Masson, Paris
Château Emile (1912) Les associations végétales. Bulletin de la Société d’Histoire Naturelle
d’Autun, pp 175–192
Clos Dominique (1864) Coup d’oeil sur la végétation de la partie septentrionale du département de l’Aude.
Mémoires de l’Académie des sciences, inscriptions et belles-lettres de Toulouse, pp 421–422
Clos Dominique (1870) Mémoires de la société des arts et des sciences de Carcassonne, p 377
Corbin A (1992) Paris-province. In: Pierre N (ed) Les lieux de mémoire 3. Les France. Traditions,
vol 2. Gallimard, Paris, pp 793–794
Coste AH (1901–1906) Flore descriptive et illustrée de la France. Paul Klincksieck, Paris
Cowles HC (1899) The ecological relations of the vegetation on the sand dunes of lake Michigan. Chicago
Dagognet François (1973) Des révolutions vertes, histoire et principes de l’agronomie. Paris, pp 51–52
de Jussieu A-L (1789) Genera Plantarum. Herissant, Paris
Drouin J-M (1991) Réinventer la nature. L’écologie et son histoire. Desclée de Brouwer, Paris
Dupuis C (1979) Histoire naturelle et naturalisme dans la France de 1904, année de la fondation des natu-
raliste parisiens. In: Bulletin des naturalises parisiens (ed) Cahiers des naturalistes 35:69–106
Fischer J-L (1997) In: Ambrière M (ed) Stations maritimes, Dictionnaire du 19e siècle européen.
PUF, Paris, pp 1128–1129
Flahault C (1901) La flore et la végétation de la France. In: Coste Abbé Hippolyte (ed) Flore
descriptive et illustrée de la France. Paul Klincksieck, Paris, pp 3–4, 1901–1906
Fox R, Weiss G (1980) The organization of science and technology in France, 1808–1914.
Cambridge-University Press, Maison des Sciences de l’Homme, Cambridge, Paris
Gadeceau È (1909) Le Lac de Grand-Lieu. Monographie phytogéographique. Nantes
Gautier G (1898) Catalogue raisonné de la flore des Pyrénées-Orientales société agricole, scientifique
et littéraire des Pyrénées-Orientales
Gazel Larambergues Dissiton de (1862) Essai d’une géographie botanique du Tarn. Société
littéraire et scientifique de Castres, pp 317–327; 403–414
Grisebach AHR (1838) Über den Einfluss des Climas auf die Begrenzung der Natürlichen Floren.
Linnaea 12:159–200
306 P. Matagne
Laissus Y (1976) Les sociétés savantes et l’avancement des sciences naturelles. Les musées
d’histoire naturelle, 100e Congrès national des sociétés savantes, Paris, 1975, Section histoire
moderne et contemporaine et histoire des sciences. Paris, pp 41–68
Lassimonne SE (1892) Principes de topographie botanique. Durond, Moulins
Linnaeus C (1737) Genera Plantarum. Conrad Wishoff, Leiden
Lecoq H, Lamotte M (1847) Catalogue raisonné des plantes vasculaires du Plateau central de la
France. Annales scientifiques, industrielles et statistiques de l’Auvergne 9
Lecoq H (1854) Etude sur la géographie botanique de l’Europe et en particulier sur la végétation
du Plateau central de la France.Paris pp X et p 134
Lloyd J (1844) Flore de la Loire-Inférieure. Prosper Sebire, Nantes
Lloyd J (1854) Flore de l’Ouest de la France.(5. ed. 1897) – Nantes: Èmile Gedeceau
Luquet A (1937) Recherches sur la géographie botanique du Massif Central. Revue de géographie
alpine 26(2):467–470
Loret H and Barrandon A (1876) Flore de Montpellier ou analyse descriptive des plantes vascu-
laires de l’Hérault. Montpellier
Margalef R (1968) Perspectives in ecological theory. University of Chicago Press, Chicago
Matagne P (1988) Racines et extension d’une curiosité: la Société botanique des Deux-Sèvres,
1888–1915. In: Corbin Alain (ed) Mémoire de maîtrise d’histoire contemporaine, Tours
Matagne P (1990) De la taxinomie à la phytosociologie: Eugène Simon à la Société Botanique des
Deux-Sèvres (1898–1915). Mémoire de DEA
Matagne P (1992) La tradition des jardins et la culture régionale: le cas des Deux-Sèvres de la fin
du 18e siècle à la première guerre mondiale. Bulletin de la société botanique de France 139.
Lettres Botaniques 1:5–13
Matagne P (1996) Les naturalistes au laboratoire. Bulletin d’histoire et d’épistémologie des sci-
ences de la vie 3:30–41
Matagne P (1997a) La botanique dans le Centre-Ouest (1800 à 1915). Bulletin de la Société his-
torique et scientifique des Deux-Sèvres, 3e série, Tome 5, 1e semestre: 125–247
Matagne P (1997b) Les mécanismes de diffusion de l’écologie en France, de la Révolution fran-
çaise à la première guerre mondiale. Editions du Septentrion, Villeneuve-d’Ascq
Matagne P (1998) The taxonomy and nomenclature of plant groups. In: Acot P (ed) The European
Origins of Scientific Ecology, Editions des archives contemporaines, vol 2. Gorden & Breach,
Amsterdam, pp 427–519
Matagne P (1999a) Aux origines de l’écologie. Les naturalistes en France de 1800 à 1914. Paris:
Comité des travaux historiques et scientifiques – histoire des sciences et des techniques
Matagne P (1999b) Des jardins écoles aux jardins écologiques. In: Fischer Jean-Louis (ed) Le jardin
entre science et représentation. Comité des travaux historiques et scientifiques, pp 307–315
MacMillan C (1897) Observations on the distribution of plants along shore at Lake of the Woods.
Minnesota Botanical Studies, Minneapolis
Odum EP, Odum HT (1953) Fundamentals of ecology. W. B. Saunders, Philadelphia
Pavillard J (1926) Etudes phytosociologiques en Auvergne. Clermont-Ferrand
Revol J (1910) Catalogue des plantes vasculaires du département de l’Ardèche. Paul Klincksieck, Paris
Schimper AFW (1898). Pflanzengeographie auf physiologischer Grundlage. Jena
Souché B (1901) Flore du Haut-Poitou. Clouzeau, Niort
Tournefort de JP (1694) Éléments de botanique ou Méthode pour connaître les plantes. Paris
von Humboldt A (1805) Essai sur la géographie des plantes, accompagné d’un tableau physique
des régions équinoxiales. Schoell, Paris
von Marilaun Anton Kerner (1863) Das Pflanzenleben der Donauländer. In: The background of
plant ecology. The plants of the Danube Basin. (Reprint 1977). Arno Press, New York
Warming E (1895) Plantesamfund, Grundträk af den Ökologiske Pflanzengeografi. Kopenhagen
Warming E (1896) Lehrbuch der ökologischen Pflanzengeographie, Eine Einführung in die
Kenntnis der Pflanzenvereine. Berlin
Warming E (1909) Oecology of plants. An introduction to the study of plant-communities
(expanded translation). Clarendon Press, Oxford
307
Chapter 22
Early History of Ecology in Spain, 1868–1936
Santos Casado
A. Schwarz and K. Jax (eds.), Ecology Revisited: Reflecting on Concepts, Advancing Science,
DOI 10.1007/978-90-481-9744-6_22, © Springer Science+Business Media B.V. 2011
Introduction
The early history of ecology in Spain provides an eloquent illustration of the con-
tradictory relationships between the emerging science of ecology and the tradition
of natural history that preceded it. For in Spain, as in almost every other western
country, the nascent discipline of ecology was undoubtedly grounded in the pre-
existing knowledge, practices and institutional framework of natural history.
However, in the comparatively small and underdeveloped turn-of-the-century
Spanish scientific community, those who strove to develop aquatic ecology or plant
ecology as new, differentiated and respectable disciplines found little support
among their colleagues and were seldom paid any attention by official authorities.
Borrowing from ecological and biogeographical parlance, one could say that in
such a small area the ecological species was not able to find its own niche, or that
competitive pressure prevented it from establishing a viable population.
This, in a nutshell, is the story that is to be told in this paper. A more detailed,
fine-grained account of the reception and early development of ecology in Spain
can be found in the book “Los primeros pasos de la ecología en España” (Casado
1997), which covers roughly the same period to be examined here, from the 1868
democratic revolution to the onset of the Civil War in 1936.
Young Darwinians
A detailed account of the institutional structure of Spanish science during the late
nineteenth and early twentieth centuries is far beyond the scope of this study. A
brief summary of the knowledge available on this topic (Sánchez Ron 1999; López-
Ocón Cabrera 2003) should be enough, however, to highlight some characteristics
of the institutionalization process relevant for further discussion.
S. Casado (*)
Universidad Autónoma de Madrid, Spain
e-mail: santos.casado@uam.es
308 S. Casado
The nineteenth century was one of decline for Spain, which lost most of its
colonies in America, and became a peripheral power in Europe. The internal political
situation was dominated by social and ideological conflicts, which were met with
repressive policies on the part of the absolutist and conservative forces that, with
few exceptions, ruled the country. In this somewhat unfavourable context, it was
left to several “intermediate” generations of Spanish scientists (López Piñero 1992)
to continue pursuing research and publishing activities, thus acting as a bridge until,
in the last third of the century, the situation improved.
The 1868 democratic revolution brought ideological freedom, which favoured scien-
tific modernization, including the spread of Darwinism. It was also the starting point for
a period of institutionalization of Spanish science. Naturalists led the way, founding in
1871 the “Sociedad Española de Historia Natural” in Madrid (compare the situation in
France in this period Chap. 20). In the following years, similar initiatives were taken in
other scientific fields, such as geography, anthropology and medicine. The re-establish-
ment of the monarchy in 1875 led to some ideological persecution, with scientists such
as Laureano Calderón and Augusto González de Linares, for example, temporarily los-
ing their chairs at the University of Santiago (Cacho Viu 1962); however, it also stabi-
lised the political and social situation and thus, in the long term, benefited the
institutional development of science, which had begun during the revolutionary period.
Yet neither the “Sociedad Española de Historia Natural” nor other private associations
could support research activities in any direct way. The only official natural history
research centres that existed at that time were the “Comisión del Mapa Geológico de
España” (Commission for the Geological Map of Spain), which was mainly committed
to the exploration of mining resources, and the “Museo de Ciencias Naturales”, both
based in Madrid. The latter, founded in 1771, had experienced a profound crisis when,
in 1845, it had become a mere dependency of the “Universidad Central”. All the natural
history professors at the university were simultaneously members of the museum,
which had no other research staff (Barreiro 1992). With such limited human and mate-
rial resources it should come as no surprise that, in comparison with other western
European countries, basic knowledge about the natural history of Spanish territory
contained some enormous gaps (Casado 1994). Many taxonomic groups were almost
unknown, and what little data existed was somewhat scattered. Vast regions remained
largely unexplored from the geological, botanical and zoological points of view.
In fact, many of the most valuable contributions to Spanish geology, botany and
zoology were made during the late nineteenth century by foreign naturalists from
Northern and Central Europe, such as Édouard de Verneuil, Moritz Willkomm and
Franz Steindachner, who travelled to the Iberian peninsula, attracted by its extraor-
dinarily rich and little known natural diversity in comparison. The founders of the
“Sociedad Española de Historia Natural” were well aware of this shortcoming.
Consequently, their inaugural manifesto was an invitation to collaborate on a wide
taxonomic, geographical and descriptive work on “the natural products of the
country”1 to accomplish a body of information comparable to those already existing
in other European countries (Sociedad Española de Historia Natural 1872).
1 “las producciones naturales del país” (Sociedad Española de Historia Natural 1872, p. VI).
30922 Early History of Ecology in Spain, 1868–1936
It soon became clear that producing a natural history of Spain would not be a task
that took matter of years but would occupy decades of research, so this collective
project shaped a long-lasting scientific tradition in which several generations of
naturalists were trained and their approaches and goals defined. What role, if any,
did ecological ideas have in this context? First, it must be noted that, in spite of their
lagging behind in surveying the national territory, Spanish naturalists were not unre-
ceptive to new trends in geology and biology, and particularly to Darwinism (Núñez
1977; Glick et al. 2001). While the historical connections between Darwinism and
ecology have been interpreted in a variety of ways (Stauffer 1960; Acot 1983;
Coleman 1986), there is no doubt that in the Spanish case a positive correlation can
be found, at least on the theoretical level. Those young naturalists who led the way
to the new Darwinian faith (Sala Catalá 1981) were also the first explicitly to
demand the inclusion of ecological issues in the research agenda of Spanish bota-
nists and zoologists. The following examples serve to substantiate this claim.
Ignacio Bolívar (1850–1944) was the most prominent Spanish specialist in the
taxonomic study of Orthoptera and as such was a contributor to the collective proj-
ect aimed at establishing a basic body of knowledge about Spanish nature. However,
in his first important work, a fauna of Iberian Orthoptera published when he was 25
years old, he cautioned his colleagues not to forget that the ultimate purpose of
natural history was to unveil such general problems as the interrelationships
between organisms and the environment, a protoecological issue typical of natural-
ists influenced by Darwinian theories.
Finalmente, no debo terminar sin recordar á los jóvenes entomólogos, á quienes principalmente
van dirigidas estas notas, que si es importante enumerar y dar á conocer las especies, sobre todo
en un país como el nuestro, en que tanto hay por conocer, á pesar de la incesante actividad y de
los laboriosos esfuerzos de varios naturalistas que á su estudio desde largo tiempo se dedican,
no lo es menos tratar de investigar las relaciones que entre sí y con los medios exteriores guar-
dan las diferentes especies, para indagar y dar solucion en lo posible á los grandes problemas
de la naturaleza, fin último á que tiende nuestra hermosa ciencia2 (Bolívar 1876, p. 86).
Odón de Buen (1863–1945) was one of Bolívar’s students and he readily applied
the same point of view in his first scientific paper, a survey of the flora of the
steppe-like central regions of Spain, published when he was barely 20 years old.
Instead of providing a conventional floristic catalogue, Buen attempted to elaborate
a general phytogeographical study, characterizing several plant groupings accord-
ing to different topographic and climatic conditions. The motivation for such an
approach was explained clearly as follows:
Los estudios geografico-botánicos han adquirido verdadera preponderancia. Iniciados por
Humboldt y secundados por Schouw, A. De Candolle, Wahlenberg y algunos otros, recibi-
eron considerable impulso con la aparicion de la teoría de Darwin en el horizonte de las
2 “Finally, I should not finish without reminding young entomologists, to whom these notes are
addressed in the first instance, that even if it is important to enumerate and publicize the existence
of all the species, especially in a country such as ours where so much is still unknown despite the
restless activity and laborious efforts of several naturalists long devoted to its study, it is no less
important to try to investigate the relationships that the different species have among themselves
and with their external media, in order to search for and, as far as possible, to provide solutions to
the great problems of nature, the ultimate goal towards which our beautiful science is directed”.
310 S. Casado
ciencias naturales. Buscando hechos en apoyo de las tendencias evolucionistas primero, y
alentados con el triunfo despues, diferentes sabios estudiaron las relaciones entre la planta,
el suelo que habita y el clima en que vive3 (de Buen 1883, p. 421).
Around that time, José Gogorza, also a student and colleague of Bolívar’s, was one
of the Spanish naturalists who journeyed to the famous Naples Stazione Zoologica
(Fantini 2000), a mecca for converts to the new biology. There he chose to study
the response of marine animals to freshwater conditions, a physiological and eco-
logical problem closely connected to the concept of adaptation (Gogorza 1891).
Both Buen and Gogorza were members of the “Sociedad Linneana Matritense”
(Linnean Society of Madrid). This botanical association, which brought together
mainly young naturalists, had been founded in 1878 by Tomás Andrés y Tubilla
(1859–1882). One of the initial purposes of the society was to complete an inventory
of Spanish flora; however, Andrés y Tubilla soon proposed a new project aimed at
studying the plant geography of the Iberian peninsula. He led this biogeographical turn
by proposing a new botanical division of the Iberian peninsula based on the distribu-
tion of plant species within two families (Andrés y Tubilla and Lázaro e Ibiza 1882).
Andrés y Tubilla’s biogeographical paper was published just after his death,
when he was only 22 years old. We do not know if he would have developed his
ecological interests further. What we do know is that other naturalists who remained
active – such as Bolívar and Gogorza – did not. It seems that they lost their enthu-
siasm for biogeographical and ecological subjects and soon joined the same kind of
taxonomic research programmes that their predecessors had undertaken, that is, a
mere cataloguing of Iberian flora and fauna. Thus, the initial reception of ecological
approaches among Spanish naturalists was very limited and lacked continuity. It
corresponded almost exactly with the formative years of the generation of natural-
ists born in the 1850s. From about 1890 onwards they apparently lost what could
be called their ecologically-minded scientific personality and their work was some-
how diluted in the collective project aimed at the completion of a natural history of
Spain comparable to those already available in other European countries.
Many years later, Ignacio Bolívar, the central figure of that group, was to
describe the situation in retrospect using rather telling words.
Nuestros naturalistas se han ocupado principalmente, casi exclusivamente, en el estudio de
las especies que viven en la Península, por el que necesariamente se había de empezar, pues
aparte de que en su tiempo otras modalidades de la ciencia eran desconocidas o poco
estudiadas, se imponía con toda urgencia hacer, por decirlo así, el inventario de los seres
que pueblan nuestro suelo, como base de todo estudio ulterior4 (Bolívar 1922, p. 65).
3 “Plant geography studies have become extremely important. First Humboldt and later Schouw,
A. de Candolle, Wahlenberg and a number of others led the way. Later, these studies were given
a significant boost with the appearance of Darwin’s theory in the realm of the natural sciences.
Searching initially for facts in support of evolutionary theory, and then stimulated by their success
in doing so, several scholars studied the relationships between plants and the soil and climate in
which they live”.
4 “Our naturalists have devoted themselves mainly – indeed almost exclusively – to the study of
the species that inhabit the [Iberian] Peninsula. This was the principal task that needed to be car-
ried out because, apart from the fact that other scientific modalities were unknown or little pursued
at that time, it was utterly imperative to compile an inventory of the natural beings that live in our
land, as a basis for further study”.
31122 Early History of Ecology in Spain, 1868–1936
Certainly ecology was among those “other scientific modalities” that were “unknown
or little pursued” because of Spanish naturalists’ almost exclusive dedication to the
descriptive and taxonomic “study of the species”, and so it remained for a long
time. Nevertheless, in the period between 1910 and 1930 a few scientists ran the
risk of placing themselves outside the mainstream of Spanish natural history and
launched new scientific programmes positioned firmly in the field of ecology.
Celso Arévalo and Aquatic Ecology
In 1912 Celso Arévalo (1885–1944) founded the first centre devoted to the ecologi-
cal study of continental waters in Spain in Valencia, called the “Laboratorio de
Hidrobiología Española”. Both on account of this institutional success as well as his
research and popularizing works, Arévalo must be considered to be the one Spanish
scientist who was responsible for establishing that form of aquatic ecology known
as limnology or, as Arévalo preferred, hydrobiology (Casado 1997, pp. 155–263).
Having graduated in the Natural Sciences, Arévalo undertook further research
training in 1905 at the “Estación de Biología Marítima” in Santander, which had
been founded in 1886 and at that time was the only institution involved in marine
biological studies in Spain (Fig. 22.1). Although the ecological approach did not
figure in Santander, Arévalo adopted such an approach when he founded the
“Laboratorio de Hidrobiología Española” a few years later. Arévalo took the insti-
tutional model of the marine laboratories and transferred it to continental waters,
while simultaneously shifting towards a different, more ecological orientation in
Fig. 22.1 Celso Arévalo (right) and an unidentified researcher at the “Estación de Biología
Marítima” in Santander, circa 1905 (Courtesy of María Teresa Arevalo, Madrid)
312 S. Casado
aquatic biology, in imitation of the emerging limnological schools of Central
Europe and North America. In Arévalo’s own words, it was a matter of establishing
“la Hidrobiología” in Spain “as a Science created by the new orientations of
Natural History”5, replacing the “criterio taxonómico” with what he called the new
“criterio biológico”6. The aim of the latter was to focus not on the species but on
the “grupo biológico”, defined by its association to one particular “medio”
(medium) (Arévalo 1914a).
In addition to the laboratory in Santander, new marine laboratories were
founded between 1905 and 1915 in several locations along the Spanish coast.
Arévalo argued that similar institutions should exist to study rivers and lakes, and
he attempted to attract the government’s attention to his project by emphasizing
potential applications with regard to the improvement of fishing and other eco-
nomic activities. He also argued that other European countries had already applied
the model of coastal laboratories to inland waters, building limnological research
centres in different lakes and rivers. One such example was the pathbreaking
Biological Station at Plön, in Germany (Overbeck 1989), cited by Arévalo as an
outstanding example among many others in Europe and the United States (Arévalo
1914b).
Arévalo’s professional and institutional standing was not unproblematic, how-
ever. He taught natural history in the public educational system, taking up a post at
the secondary school in Valencia, the “Instituto General y Técnico de Valencia”, in
1912 (Fig. 22.2). It seems that the proximity to the school of a coastal lagoon named
“L’Albufera” prompted Arévalo to start his project in imitation of foreign limno-
logical centres. Like those centres, Arévalo’s laboratory was set up specifically to
study a single system, L’Albufera. The laboratory itself, however, was located not
on the shore of the lagoon but a few miles away, in the city of Valencia. It had no
permanent staff, no boats and no other equipment that would have been needed to
carry out proper limnological surveys. In fact, in its early days, the “Laboratorio de
Hidrobiología” was simply a personal initiative on the part of Arévalo that had no
support except for that of the “Instituto General y Técnico de Valencia” where he
taught natural history. Arévalo had obtained permission from the head of the school
to use a free space formerly used as a corridor, where he managed to install a few
research tables with microscopes and other research equipment, as well as some
aquariums for the observation of live organisms. Only after several years of work,
in 1917, was it officially recognized by the government as a research centre. This
was largely a matter of status, because even then the laboratory continued to operate
with almost no funding from any local or national authority.
5 “como una Ciencia creada por las nuevas orientaciones de la Historia Natural” (Arévalo 1914a).
6 By “biological criterion” Arévalo simply meant that he wanted to study actual biological com-
munities rather than artificially-defined taxonomic groups. Of course, such approach was hardly
new in the 1910s, but it must be noted that these kinds of ecological studies were almost inexistent
in Spain at that time.
31322 Early History of Ecology in Spain, 1868–1936
In spite of these limitations Arévalo managed to carry on doing remarkable
research on the limnology of L’Albufera and nearby wetlands. However, soon
enough he was forced to face a paradox. Having declared that it was necessary to
change the “taxonomic criterion” dominant in Spanish natural history, he was still
confronted with the very same problem – insufficient taxonomic knowledge – that
had prompted a taxonomic-centered tradition among Spanish naturalists. The short-
coming was even worse for aquatic groups, which had been especially neglected.
As a consequence, Arévalo was forced to begin his work by making taxonomic
surveys of the aquatic organisms that were of interest to him, especially the plank-
tonic groups. This contradictory situation prevented a deeper development of
Arévalo’s ecological research. However, the difference in his approach was clear.
Instead of trying to specialize in one or a few groups for the whole Iberian penin-
sula, Arévalo concentrated on L’Albufera, studying as many groups as possible.
This is how he became the first Spanish scientist to publish works on such impor-
tant groups as Cladocera and Rotatoria (Arévalo 1916, 1917). In addition, he pro-
duced many short notes and gathered unpublished data on almost all the aquatic
groups that inhabited L’Albufera, from algae to birds. Arévalo also investigated the
spatial distribution of organisms in relation to the physicochemical conditions of the
Fig. 22.2 An employee of the “Instituto General y Técnico de Valencia” shows part of the
limnological equipment of the “Laboratorio de Hidrobiología”, circa 1915 (Courtesy of María
Teresa Arévalo, Madrid)
314 S. Casado
lagoon, and became especially interested in the temporal variation, both qualitative
and quantitative, of planktonic communities. However, it seems that he did not
attempt to study interspecific relationships.
Arévalo was a member of the “Sociedad Española de Historia Natural”, where
he talked to his colleagues about the new laboratory and its scientific goals (Arévalo
1914b), but his ecological projects did not enjoy much support from the scientific
establishment in Madrid. In order to seek more local support, he founded a branch
of the society in Valencia, becoming the leader of a small group of Valencian non-
professional naturalists who provided some practical assistance for his project.
Arévalo’s main follower and collaborator was biologist Luis Pardo (1897–1957).
Some foreign naturalists were temporarily affiliated with the laboratory, which
proved useful for their specialized studies on aquatic groups. Such was the case
with German malacologist Fritz Haas, German specialist on water mites Karl Viets,
and Swiss ichtyologist Alfonso Gandolfi. Arévalo even managed to create a regular
forum for his own publications by creating a journal for the Instituto, the “Anales
del Instituto General y Técnico de Valencia”. This contained a special series enti-
tled “Trabajos del Laboratorio de Hidrobiología Española” which included 33
limnological works from 1916 to 1928. The “Trabajos” were reprinted separately
for the purpose of scholarly exchange with foreign limnological journals and cen-
tres. It should be noted that with regard to limnology, most of Arévalo’s scientific
relationships were with foreign researchers and centres, especially in German-
speaking countries. In 1921 he travelled to France, Switzerland and Germany to
visit several limnological laboratories, including the one at Kastanienbaum, near
the Swiss city of Lucerne, which was directed by Hans Bachmann. He also became
a member of the “International Association of Theoretical and Applied Limnology”
(best known as SIL) immediately after its founding in 1922, and attended the SIL
congress in Rome in 1927.
In summary, Arévalo created his own institutional framework for his ecological
project, taking an alternative route to the taxonomic approach favoured by the offi-
cial centres devoted to natural history.
In 1919 Arévalo took up a new post to teach natural history at the “Instituto del
Cardenal Cisneros” in Madrid. Subsequently, the “Museo Nacional de Ciencias
Naturales” in Madrid created a new section of hydrobiology for Arévalo, who was
appointed head of department. The “Laboratorio de Hidrobiología Española” in
Valencia was incorporated as a branch of the museum. Did this mean that the
museum was suddenly interested in Arévalo’s ecological project? Clearly not – in
fact, the museum’s interest lay in transforming the laboratory in Valencia into a
marine station. The museum had lost control over several coastal stations due to a
previous institutional reorganization of those scientific centres controlled by the
government, and it was in urgent need of alternative seashore locations if it was to
carry on the taxonomic studies of marine groups that were the subject of several of
its staff’s zoologists. Nonetheless, this agreement gave Arévalo the chance to main-
tain some control over his laboratory in Valencia, which would have probably been
lost otherwise; most importantly, it gave him the official support that he had sought
for so long.
31522 Early History of Ecology in Spain, 1868–1936
It was difficult for both the museum and Arévalo to safeguard their own respec-
tive interests. After some time the museum decided that Valencia was not the right
place to establish a coastal station and lost all interest in the laboratory. In the
meantime, Arévalo had succeeded in establishing an official post in Valencia for his
collaborator Luis Pardo, who was appointed assistant researcher at the laboratory.
For a few years Arévalo kept up his limnological research, focusing on the temporal
variation of plankton and surveying various Spanish lakes, sometimes in collabora-
tion with Pardo. It seems that this kind of research was not held in high regard by
the museum’s director, Ignacio Bolívar, or by other influential members of the
centre, and soon the support given to Arévalo was reduced to a minimum. As men-
tioned in the previous section, Bolívar had long lost his youthful enthusiasm for
protoecological subjects by this time. In 1928 Pardo moved to Madrid to attend to
personal business and applied, without success, to continue his work as Arévalo’s
assistant at the museum in Madrid. In December 1931 Arévalo resigned his post at
the museum. The laboratories of hydrobiology in Madrid and Valencia were finally
closed down in 1932.
During his final years at the “Museo Nacional de Ciencias Naturales” Arévalo
had sought alternative sources of support for his project. Instead of seeking help
from scientific institutions controlled by naturalists, he addressed himself to gov-
ernment agencies dealing with natural resources of economic interest. He rein-
vented himself as a specialist in applied biology and drew attention to the state
of neglect suffered by inland fisheries up to this point. This professional
transformation was not Arévalo’s preferred path – “[Y]ou know that I am not
interested in technical aspects of fishing, but only in those that are purely scien-
tific”7 said Arévalo in a letter to Pardo in 1927. However, it proved to be the only
effective way to institutionalize limnological studies in Spain. He also played an
active role in various official commissions appointed to reform the legal regulation
of fishing, insisting on the importance of biological studies as the only sound basis
for improving the economic and social outputs of continental waters.
Management of inland fisheries was one of the tasks officially assigned to the
corps of forestry engineers. As a distinct professional branch in the Spanish
governmental administration, forestry engineers had their own official centre for
applied research, the “Instituto Forestal de Investigaciones y Experiencias” (Institute
of Applied Forestry Research). In 1931 this centre created a new “Section of
Biology of Continental Waters”. Thanks to the contacts made by Arévalo in previ-
ous years Pardo became a member of the technical staff of this new department.
Arévalo’s own scientific career in aquatic ecology was virtually finished by
1931. But even after his death in 1944, Arévalo’s legacy remained alive in his book
“La vida en las aguas dulces”, which, in spite of its popularizing character, gave a
remarkably complete and updated view of the field and must be regarded as the first
treatise on limnology published in Spanish (Arévalo 1929).
7 “[Y]a sabes que yo no me intereso por los problemas tecnicos de la pesca y solamente por los
puramente cientificos” (Casado 1997, p. 240).
316 S. Casado
An offshoot of Arévalo’s project was Luis Pardo’s affiliation with the “Instituto
Forestal”, where the latter continued studying Spanish continental waters, even if
for the most part his focus was now on applied aspects, especially fishing activities.
Apart from being Arévalo’s collaborator, Pardo was significant because he was the
first to attempt a comprehensive survey of Spain’s rivers and lakes as natural
resources. Even so, pure ecology was also represented at the “Instituto Forestal”, as
exemplified in the remarkable monograph on the river Manzanares written by for-
estry engineers Luis Vélaz de Medrano and Jesús Ugarte, which was the first eco-
logical study of riverine ecosystems to be carried out in Spain (Vélaz de Medrano
and Ugarte 1934). However, all opportunity to continue aquatic ecological research
was eliminated by the outbreak of the Spanish Civil War in 1936. That is why, after
the war, the young naturalist Ramon Margalef, unknown at that time, had to start
his career in aquatic ecology from scratch. Ramon Margalef was to become one of
the world’s most influential ecologists in the 1960s and the 1970s – but that is a
different story.
Emilio H. Del Villar and Plant Ecology
While Arévalo developed his project in the field of aquatic ecology, another
Spanish naturalist chose plant ecology as his own scientific project. Emilio Huguet
del Villar (1871–1951) is arguably the most influential of the scientists who under-
took ecological research in Spain before 1936 (Casado 1997, pp. 265–352). He was
the only one to gather around him a core group of researchers that might have
become a genuine ecological school. He was also the most ambitious in defining
his research programme, which was aimed at producing a complete, ecologically-
based update of the plant geography of the Iberian peninsula. Finally, he was note-
worthy for the originality of his theoretical elaborations, many of which he
summarized in his book “Geobotánica” (del Villar 1929). But again, Villar’s eco-
logical project found little support among Spanish naturalists and he was not able
to create a stable institutional structure for it.
Villar was a self-taught scientist without a university degree. This may well have
been a key factor in the originality of his scientific thought. Villar’s choice of plant
ecology – a field almost completely ignored in Spain at that time – and the innova-
tive way in which he developed it are characteristic of his overall personality.
However, lacking any official academic qualification proved to be an additional
obstacle in his somewhat eccentric career: having started out in journalism, he turned
first to geography and later to botany and plant ecology, eventually ending up in the
field of soil science (Martí 1984). From both an intellectual and institutional point
of view, Villar is a prime example of the “scientific outsider”, a characterization
I have applied to early Spanish ecologists elsewhere (Casado 1997, p. 442).
Villar began his autodidactic training as a botanist and plant ecologist around
1912, but, lacking any professional links with scientific institutions, he earned his
living writing geography books intended for a broad audience. In fact, the work that
31722 Early History of Ecology in Spain, 1868–1936
may be considered his first publication on the ecology of Iberian vegetation appeared
in 1921 as part of one of these geographical books (del Villar 1921, pp. 176–192).
Four years later Villar published a much more detailed report, entitled “Avance geo-
botánico sobre la pretendida estepa central de España” (Preliminary geobotanical
study on the presumed central steppe of Spain) (del Villar 1925). For this paper,
which was to be his official presentation as a fully-fledged ecologist, Villar cleverly
chose a subject which demonstrated that modern ecological theories and methods
could produce a radically different interpretation of vegetation. Adopting a dynamic
approach which, by his own admission, was closely modelled on Frederic E.
Clements’ successional plant ecology, Villar showed that what had hitherto been
known as the Spanish “central steppe” was in fact the result of a long process of
anthropogenic deforestation and that the remnants of the original forests could be
used to reconstruct the composition and structure of the climax vegetation.
During this period Villar had been engaged professionally as a plant ecologist,
for the first and only time in his life, in his native Catalonia. In 1923 botanist Pius
Font i Quer, who was serving as director of the “Museu de Ciències Naturals de
Barcelona” at that time, had offered Villar a post in a new section of the museum
devoted to plant geography. This museum was one of the scientific institutions that
had been created during the first decades of the twentieth century as part of a
broader cultural drive initiated by the Catalan nationalist movement. They repre-
sented an alternative to the central institutions based in Madrid, where Villar had
found little support. Villar accepted this post but, wanting to establish his own
research programme, he soon came into conflict with the museum and was dis-
missed in 1924. Villar had been a member of the Madrid-based “Sociedad Española
de Historia Natural”, where he published his first botanical works, since 1915, but
his work attracted little attention. In 1919 he joined a new association, the “Sociedad
Ibérica de Ciencias Naturales” (Iberian Society of Natural Sciences), based in
Saragossa. A year later Villar founded a branch of this society in Madrid, clearly
seeking an alternative scientific audience for his proposals and findings.
The quality and originality of Villar’s work did not go completely unnoticed
among Spanish botanists, especially the younger ones. The publication of his book
“Geobotánica” (Villar 1929), the first treatise on plant ecology to be published in
Spanish, further enhanced his prominent status by offering a concise and up-to-date
summary of the field (Fig. 22.3). In addition, it proposed a large number of theoretical
innovations, such as his classification of vegetation types and his concept of “sine-
cia”, which Villar intended to introduce as a bias-free term for any unit of vegetation,
equally distant from the sociologically-laden terms “community” and “association”.
These innovations endowed his work with the nationalist appeal of a genuinely
Spanish contribution to vegetation science. Soon several university professors, gradu-
ate students and amateur botanists were following Villar’s ideas and methodology.
One of them was José Cuatrecasas (1903–1996), professor of Botany at the
“Universidad Central” of Madrid, who combined his dedication as a plant taxonomist
specializing in tropical flora with regional ecological studies of Iberian vegetation,
which he carried out in close cooperation with Villar. This might have become the
starting point for a Spanish school of plant ecology had Villar managed to pursue his
318 S. Casado
ecological interests further. Of course the term “school” is used here very loosely, for
almost none of the conditions that have been proposed to define a research school
(Morrell 1972; Geison 1981; Servos 1993) were met by the informal and somehow
blurry group of naturalists clustered around a scientist without any institutional affili-
ation as Villar. In fact, the professional standing of Villar’s followers was often much
better than his own.
Such an anomalous situation may partially explain a new shift in Villar’s scien-
tific career towards soil science, which can be dated at about 1925. Villar started out
with an interest in soils as one of the key factors in plant ecology. As soil science
was almost unknown in Spain, he sought advice from leading specialists abroad and
in 1924 attended the International Conference of Soil Science held in Rome. The
International Association of Soil Science was created at the Rome conference and
Villar was invited to organize the Spanish section. For Villar this was an excellent
opportunity to ask for official support. Soil science, Villar argued, had obvious appli-
cations and potential economic benefits that the Spanish government could no longer
afford to ignore. In 1925 an official commission was created to serve as the Spanish
section of the International Association of Soil Science. Villar was appointed execu-
tive secretary and he even managed to include plant ecology – still his main interest
– among the subjects covered by the new “Comisión de Edafología y Geobotánica”
(Commission for Soil Science and Geobotany). But soil science was the commis-
sion’s priority and Villar, who was the only researcher among its members, had to
devote himself to these kinds of studies, largely abandoning his investigations in
plant ecology. Once again, forestry engineers provided the institutional support that
Fig. 22.3 Classification of vegetation types proposed by Emilio H. del Villar in his book
“Geobotánica” (Villar 1929)
31922 Early History of Ecology in Spain, 1868–1936
naturalists had been reluctant to give. From 1927 to 1932 Villar worked at the “Soils
Section” at the “Instituto Forestal de Investigaciones y Experiencias”. Villar is
generally credited as being the founder of soil science in Spain, and he eventually
published the first map of soils of the Iberian peninsula (Martí 1984).
Spanish botanists with an interest in ecology were “orphaned”, as it were, by
Villar’s progressive withdrawal from ecology, and therefore turned their attention to
Montpellier in France, where Josias Braun-Blanquet’s phytosociological school had
its headquarters at the “Station International de Géobotanique Méditerranéenne et
Alpine”, best known as Sigma (see also Chap. 21). Sigma had been founded by
Braun-Blanquet in 1930 and was especially influential as an international training
centre (Acot 1993). It was therefore an obvious choice for those wishing to specialize
in vegetation research. In 1934 three young Spanish botanists attended courses at
Sigma, and that same year an international excursion organized by Sigma and led by
Braun-Blanquet visited Catalonia. José Cuatrecasas helped to organise these activi-
ties, which were intended to introduce the so-called Sigmatist school into Spain
(Fig. 22.4). While Cuatrecasas himself was an adherent of Villar’s concepts and
methods, he understood that Sigma’s structure could be used to fill the institutional
gap that was preventing further development of plant ecology in Spain. There were
profound differences, both in theory and practice (Nicolson 1989), between the taxo-
nomic, static approach of Sigmatist phytosociology and the more holistic, dynamic
approach of Clementsian plant ecology preferred by Villar. Phytosociology was con-
cerned with floristic inventories and with the classification of vegetation types; Villar’s
plant ecology, by contrast, focused on the relationships between plant communities
Fig. 22.4 José Cuatrecasas (left), a local guide, and botanist Daniel Sans (right) at Sierra Magina
(Jaén, Spain), where Cuatrecasas initiated his ecological investigations in the early 1920s
(Courtesy of the Institut Botànic de Barcelona)
320 S. Casado
and environmental factors with the aim of producing a successional interpretation of
vegetation. Nevertheless in the peculiar Spanish context of the 1930s, these influ-
ences should be considered as complementary rather than in opposition. Villar’s
works had aroused interest in plant ecology among Spanish botanists, and Sigma
provided a methodological and institutional framework.
The outbreak of the Civil War in 1936 threw the scientific community into com-
plete disarray and interrupted the ongoing acclimatization of plant ecology among
Spanish naturalists. Both Villar and Cuatrecasas suffered the ideological persecu-
tion that followed the victory of General Franco, which forced many Spaniards into
exile. Thanks to the international prestige he had achieved as a soil scientist, Villar
was able to establish himself in Algeria and, later, in Morocco, where he continued
studying soils until his death in 1951. Cuatrecasas settled in Colombia and later in
the United States, where he continued his taxonomic studies as a specialist in tropical
flora. He died in Washington DC in 1996.
Meanwhile, a new generation of botanists was in the process of filling the pro-
fessional gap left in Spain by “émigrés” after the war. Initially, some of them super-
ficially adopted Villar’s methodology, but Sigmatist phytosociology was soon
reintroduced by botanists such as Salvador Rivas Goday (Izco 1981) and Oriol de
Bolòs (Pairolí 2001) – this time in an overwhelmingly dominant way, which has
had a profound influence on vegetation research in Spain ever since.
Scientific Outsiders and Interrupted Projects
In some ways both Arévalo and Villar were “scientific outsiders” (Casado 1997, p.
442). They never held a university chair or a post in a pre-existing research depart-
ment. They chose, instead, to create their own intellectual and professional oppor-
tunities, identifying previously unoccupied niches – that is to say, disciplines which
had received little or no attention by Spanish naturalists – such as aquatic and plant
ecology. They enjoyed more support – at least temporarily – in peripheral cities
such as Valencia and Barcelona, which had more dynamic and open institutional
settings, than in Madrid, where the central, traditional scientific institutions were
based. Their strategy enjoyed mixed fortunes. On the one hand, they succeeded in
establishing themselves as authorities in their fields, and created, albeit provision-
ally, an institutional basis for their projects, such as the “Laboratorio de
Hidrobiología Española” and the “Comisión de Edafología y Geobotánica”. On the
other hand, their achievements proved to be built on sandy soil and were quickly
washed away when unfavourable personal, social or political contingencies arose.
The lack of support from the scientific establishment in the fields of natural history,
botany and zoology, where their ecological projects should have been embedded,
jeopardized the continuity of their pursuits, which they eventually abandoned.
Thus the early development of ecology in Spain was very limited and did not
play a significant role in relation to international ecology. But it provides an interesting
example of the complex processes and contradictory relationships involved in the
development of new disciplines within the traditional framework of natural history.
32122 Early History of Ecology in Spain, 1868–1936
In Spain, as in many other countries, naturalists were the natural substratum for the
reception of new ecological approaches (Kohler 2002; Kingsland 2005). However,
the fruitful synthesis envisioned in those early years by pioneer ecologists such as
Charles C. Adams – “bringing together the best of old natural history and of the
new laboratory biology” (Ilerbaig 2000, p. 459) – could not happen in Spain.
Insufficient taxonomic knowledge certainly posed a problem for Arévalo and
Villar, but this was not the real difficulty. The national and nationalist tradition that
had shaped turn-of-the-century Spanish natural history into a taxonomy-oriented
project proved to be a major obstacle for those early ecologists in terms of acquiring
a recognized institutional and professional status. Even though Arévalo and Villar
were primarily interested in basic ecological research, both of them were forced to
emphasize the applied potential of their research fields in order to obtain alternative
support from official institutions that were controlled by forestry engineers and
devoted to fisheries, forestry and agriculture.
As far as I am aware, Arévalo and Villar had no personal or professional contact
with one another, nor did they appear to be conscious of the connection between
their scientific interests and professional careers. However, their common ecological
approach can surely be linked to similarities in the historical development of their
projects, including the way they related to the intellectual and social context within
the turn-of-the-century Spanish scientific community.
Fig. 22.5 The map displays all the locations of early Spanish ecology activities mentioned in the
article (arranged by C. Haak)
322 S. Casado
References
Acot P (1983) Darwin et l’écologie. Revue d’Histoire des Sciences 36:33–48
Acot P (1993) La phytosociologie de Zürich-Montpellier dans l’écologie française de l’entre-deux
guerres. Bulletin d’ Ecologie 24:52–56
Andrés y Tubilla T, Lázaro e Ibiza B (1882) Distribución geográfica de las columníferas de la
Península Ibérica. In: Resumen de los trabajos verificados por la Sociedad Linneana Matritense
durante el año 1881. Sociedad Linneana Matritense, Madrid, pp 25–33
Arévalo C (1914a) La Hidrobiología como Ciencia creada por las nuevas orientaciones de la
Historia Natural. Ibérica 2:317–319
Arévalo C (1914b) El Laboratorio hidrobiológico del Instituto de Valencia. Boletín de la Real
Sociedad Española de Historia Natural 14:338–348
Arévalo C (1916) Introducción al estudio de los Cladóceros del plankton de la Albufera de
Valencia. Anales del Instituto General y Técnico de Valencia 1(1):1–67
Arévalo C (1917) Algunos rotíferos planktónicos de la Albufera de Valencia. Anales del Instituto
General y Técnico de Valencia 2(8):1–50
Arévalo C (1929) La vida en las aguas dulces. Labor, Barcelona
Barreiro AJ (1992) El Museo Nacional de Ciencias Naturales (1771-1935). Doce Calles, Aranjuez
Bolívar I (1876) Sinópsis de los ortópteros de España y Portugal. Anales de la Sociedad Española
de Historia Naural 5:79–130
Bolívar I (1922). Contestación. In: García Mercet R.Discurso leido en el acto de su recepción por
el Señor D. Ricardo García Mercet. Madrid: Real Academia de Ciencias Exactas, Físicas y
Naturales, pp. 49–70.
De Buen O (1883) Apuntes geográfico-botánicos sobre la zona central de la Península Ibérica.
Anales de la Sociedad Española de Historia Natural 12:421–440
Cacho Viu V (1962) La Institución Libre de Enseñanza. Rialp, Madrid
Casado S (1994) La fundación de la Sociedad Española de Historia Natural y la dimensión nacio-
nalista de la historia natural en España. Boletín de la Institución Libre de Enseñanza 19:45–64
Casado S (1997) Los primeros pasos de la ecología en España. Ministerio de Agricultura, Pesca
y Alimentación, Madrid
Coleman W (1986) Evolution into ecology? The strategy of warming’s ecological plant geogra-
phy. J Hist Biol 19:181–196
Fantini B (2000) The history of the stazion Zoologica Anton Dohrn. An outline. In: Cariello L,
Consiglio D (eds) Stazione Zoologica Anton Dohrn. Activity Report 1998/1999. ImPrint,
Napoli, pp 71–107
Geison GL (1981) Scientific change, emerging specialties, and research schools. Hist Sci
10:20–40
Glick TF, Ruiz R, Puig-Samper MA (eds) (2001) The reception of Darwinism in the Iberian world.
Kluwer Academic Publishers, Boston
Gogorza J (1891) Influencia del agua dulce en los animales marinos. Anales de la Sociedad
Española de Historia Natural 20:221–271
Ilerbaig J (2000) Allied sciences and fundamental problems: C. C. Adams and the search for
method in early American ecology. J Hist Biol 32:439–463
Izco J (1981) Prof. Salvador Rivas Goday. Lazaroa 3:5–23
Kingsland SE (2005) The evolution of American ecology. The John Hopkins University Press,
Baltimore
Kohler RE (2002) Landscapes and labscapes: exploring the lab-field border in biology. University
of Chicago Press, Chicago
López Piñero JM (1992) Introducción. In: López Piñero JM (ed) La ciencia en la España del siglo
XIX. Marcial Pons, Madrid, pp 11–18
López-Ocón Cabrera L (2003) Breve historia de la ciencia española. Alianza, Madrid
Martí J (1984) Emilio Huguet del Villar (1871-1951). Cincuenta años de lucha por la ciencia.
Universitat de Barcelona, Barcelona
32322 Early History of Ecology in Spain, 1868–1936
Morrell JB (1972) The chemist breeders: the research schools of Liebig and Thomson. Ambix
19:1–46
Nicolson M (1989) National styles, divergent classifications: a comparative case study from the
history of French and American plant ecology. Knowledge Soc 8:139–186
Núñez D (1977) El darwinismo en España. Castalia, Madrid
Overbeck J (1989) Plön history of limnology, foundation of SIL and development of a limnologi-
cal institute. In: Lampert W, Rothhaupt KO (eds) Limnology in the federal republic of
Germany. International Association for Applied and Theoretical Limnology, Plön, pp 61–65
Pairolí M (2001) Oriol de Bolòs. Una vida dedicada a la botànica. Fundació Catalana per a la
Recerca, Barcelona
Sala Catalá J (1981) El evolucionismo en la práctica científica de los biólogos españoles del siglo
XIX (1860-1907). Asclepio 33:81–125
Sánchez Ron JM (1999) Cincel, martillo y piedra. Historia de la ciencia en España (siglos XIX y
XX). Taurus, Madrid
Servos JW (1993) Research schools and their histories. Osiris 8:3–15
Sociedad Española de Historia Natural (1872) Circular. Anales de la Sociedad Española de
Historia Natural 1:5–7
Stauffer RC (1960) Ecology in the long manuscript version of Darwin’s Origin of Species and
Linnaeus’ Oeconomy of Nature. Proc Am Philos Soc 104:235–241
Vélaz de Medrano L, Ugarte J (1934) Estudio monográfico del río Manzanares. Instituto Forestal
de Investigaciones y Experiencias, Madrid
Del Villar EH (1921) El valor geográfico de España. Ensayo de Ecética. Sucesores de Rivadeneyra,
Madrid
Del Villar EH (1925) Avance geobotánico sobre la pretendida estepa central de España. Ibérica
23:281–283, 297–302, 328–333, 344–350
Del Villar EH (1929) Geobotánica. Labor, Barcelona
325
Chapter 23
Plant Community, Plantesamfund
Peder Anker
P. Anker (*)
Gallatin School of Individualized Study and Environmental Studies Program at New York
University, New York, USA
e-mail: pja7@nyu.edu
The First Use of the Plant Community Concept
The plant community concept was first introduced by the Danish botanist Johannes
Eugenius Bülow Warming (1841–1924) in his book “Plantesamfund” of 1895,
where he suggested a general theory of explaining different geographical distribu-
tions of plants. The title “Plantesamfund” can be translated both as Plant societies
and Plant communities, since the Danish word samfund means both “society” and
“community” (or alternatively “Gesellschaft” and “Gemeinschaft” in German). To
keep the broad meaning of the original title Warming chose the German title
“Ökologischen Pflanzengeographie” (1896) and the English title “Oecology of
Plants” (1909). The book addressed different factors limiting the geographical dis-
tribution of different plants. He used the concept of “community” or “Gemeinschaft”
when describing smaller geographical distributions of plants, while “oecologie” or
“Ökologie” had a broader geographical meaning corresponding to “society” or
“Gesellschaft” as a whole. “Plantesamfund” was not translated into French, though
Warming was inspired by the French botanical notion of “le commensal” (dinner
partner) in his thinking about the plant community. “Plantesamfund” was also
translated into Polish in 1900 and Russian in 1901.
Summary
The Danish botanist Warming coined the plant community concept in his book
“Plantesamfund” in 1895. It has a neo-Lamarckian, morphological, and religiously
informed understanding of plant geography. The community concept also drew its
inspiration from the Danish political and social environment. Warming was a patri-
© The U.S. Government’s right to retain a non-exclusive, royalty-free licence in and to any
copyright is acknowledged.
326 P. Anker
otic defender of the King’s council’s ambition to expand the Danish Empire and the
exploitation of natural resources. The plant community concept provided a tool for
management of nature that was inspired by the King’s steering of human communi-
ties. Warming’s morphologically informed research in Brazil and his geographical
explorations of Greenland were also of key importance in the development of his
plant community concept.
Main Phases of the History of the Concept
The plant community concept was first introduced in Danish. The English transla-
tion of 1909 is true to the original Danish definition:
The term ‘community’ implies a diversity but at the same time a certain organized unifor-
mity in the units. The units are the many individual plants that occur in every community,
whether this be a beech-forest, a meadow, or a heath. Uniformity is established when cer-
tain atmospheric, terrestrial, and any of the other factors discussed in Section I [light, heat,
humidity, air, nutrients, soil, water, etc.] are co-operating, and appears either because a
certain, defined economy makes its impress on the community as a whole, or because
a number of different growth-forms are combined to from a single aggregate which has a
definite and constant guise. (Warming 1909, p. 91).
The plant community concept emerged from Warming’s research in the Brazilian com-
munity Lagoa Santa in the early 1860s, research which was published in his book of the
same name in 1892. His voyage to Greenland in 1884 was also important to the plant
community concept, since it was during this expedition that he learned to appreciate the
importance of having a managerial overview on a geographical landscape before ana-
lyzing its plants. Warming’s patriotic political views and support of the King’s geo-
graphical ambitions for enlarging the Danish Empire were also of significance for his
plant community concept. It was assumed that only a stable and harmonious human
community could be a true resource for the nation. Plant communities, by analogy,
Warming argued, could only be a natural resource in so far as they lived in commensal-
ism with the rest of nature. He would quote the French botanist Pierre Joseph van
Beneden’s (1809–1894) definition of commensalism – “Le commensal est simplement
un compagnon de table” (The dinner partner is simply a companion at the table) – to
evoke the sense of mutually benefiting way of living he thought both humans and plants
were striving for (Warming 1909, p. 92). This commensalism was to describe a symbi-
otic relationship where different plants could live side by side at the same dinner table
without harming each other’s living conditions. He was particularly interested in cases
where certain plants may benefit from living in co-relationship with other plants.
Warming was widely read and appreciated among Danish and Scandinavian
ecologists (Prytz 1984; Söderqvist 1986). German plant geographers, such as
Andreas Franz Wilhelm Schimper (1856–1901), were also inspired by Warming,
and altogether three different versions of “Plantesamfund” appeared in the German
language (Schimper 1898; Goodland 1975).
In Britain Arthur George Tansley (1871–1955) pursued a mechanist informed
reading of Warming’s plant community concept, while his rival Isaac Bayley
32723 Plant Community, Plantesamfund
Balfour (1853–1922) pursued a morphological interpretation that was more in
accordance with Warming’s original ideas. The South African ecologist and
Balfour-student John Phillips (1899–1987) would coin the phrase “biotic commu-
nity” in reference to Warming and to the holistic philosophy of the South African
statesman Jan Christian Smuts (1870–1955). (Smuts 1926; Phillips 1931).
Warming showed little interest in epistemology and philosophy, but regarded
himself instead as a strictly empiricist. He had a religiously informed understanding
of both the human and the plant community, believing that God’s goodness and
purpose was an acting force in nature and human communities alike.
Historical Background
The plant community concept Warming developed grew out of his patriotic as well
as deeply religious views. He was raised on a farm in conservative rural Nørup west
of Velje. This landscape in Denmark was dominated by the Randbøl heath where
Warming would spend his youth nurturing a passion for nature. His father was a
priest who died when he was only three years old, and his religious views were
mixed with a lifelong longing for his father. His mother’s family consisted of
wealthy shopkeepers, and Warming would eventually inherent a fortune that
enabled him to pursue his high-society botanical interests. This he did from the age
of eighteen at the University of Copenhagen, where he would read for general
exams in natural history and botany between 1859 and 1862 (Christiansen 1924–1926,
pp. 617–665, 776–806).
This was a tumultuous political and social period in Danish history. The imperial
ambitions of the King’s cabinet caused much tension with respect to control of the
rebellious duchies of Nord Schleswig, Holstein, and Lauenburg. A bitter war to
defend the region between 1848 and 1851 did not settle the conflict, which re-
emerged in another war between 1863 and 1864 in which Prussia took control of
the duchies. In the coming years it became the King’s council’s official policy to
expand the Danish Empire as well as trying to reunite Denmark with the lost land.
Warming was on a lifelong crusade for the cause. It is telling that his biographer
describes the reunion of Nord Schleswig (now Søderjylland) with Denmark in 1920
as “the most joyful event of his life” Christiansen (1924–1926, p. 780).
His religious beliefs were, like most of his fellow Danes, Lutheran protestant.
Religion in Denmark was at the time a political matter. Though the throne provided
citizens with religious freedom and the Church was declared independent of the
State, the King’s cabinet was in reality the head of Church. In this hierarchy the
King’s good will would secure the religious purpose and order of society, commu-
nities, and the use of natural resources. Warming was a religious patriot, which
meant supporting the authority of the King, religious and social stability, and the
Danish imperial ambitions. He saw the wisdom of the King’s council in view of the
larger purpose and goodness of all living things, a purpose which had its ultimate
cause in the Creator. It was God who once started the evolution of the Creation, and
the botanists could unveil His purpose in the successive development of the living
328 P. Anker
things towards a gradually better world. It was then up to the King’s council to
wisely use botanical knowledge to guide the use of natural resources. According to
his neo-Lamarckian views, plants adapting to their environment and God’s purpose
and goodness was behind these processes in nature. Though Warming in the late
1870s adopted the Darwinian principles of evolution, he could not agree with the
view that this evolution was accidental or without a deeper aim (Coleman 1986).
This purpose in biological evolution as well as human history he understood in
view of the King’s council’s ambition of expanding the Danish Empire to secure
the wealth of the Danish nation through exploitation of natural resources.
In 1863, the same year the King’s council decided to pursue their geographical
objective of trying to take back their lost duchies with military might, Warming left
for Brazil. He was invited to serve as a secretary for the palaeontologist Peter
Wilhelm Lund (1801–1880) who was working on an excavation site in the com-
munity of Lagoa Santa. According to Warming, it was a place of “light and joy and
peace” Christiansen (1924–1926, pp. 624). It was also a place of loneliness. Lund
was a rather asocial person whose only demand of Warming was to read and orga-
nize his correspondence. This forced Warming to stay in proximity of their house,
and thus to limit his own research to the immediate surroundings. Over the next two
and a half years, he consequently came to know the geographical location of almost
every plant in the neighbourhood. Upon his return he would use three decades to
describe the fourteen cases of species he collected into a book he eventually pub-
lished as “Lagoa Santa” in 1892.
Warming was an admirer of Alexander von Humboldt (1769–1859), who in his
books about plant geography relied on morphological methods as well as system-
atic botany (Nicolson 1983, pp. 12–73). His chief source of inspiration, however,
was neo-Lamarckism and the idea that plants adapt to each other and their respec-
tive environments. Upon his return from Brazil he would plunge into morphological-
organic studies of plants. His colleagues in Denmark initially found Warming
arrogant and disagreed with his ideas about the importance of environmental factors
in understanding the geographical distribution of plants. After extensive travelling
at various universities in Germany, he eventually settled for a professorship in
Stockholm where he would lecture and write about systematic botany. These lec-
tures resulted in a series of textbooks, which were widely used in Scandinavian
universities and beyond. His “Haandbok i den systematiske botanik” (Handbook in
Systematic Botany) from 1879 and “Den almindelige botanik” (The common bot-
any) from 1880 were both reprinted in several editions as late as 1891 and 1895
respectively. When he finally received a professorship at the University of
Copenhagen in 1886 he would lecture regularly for medical and pharmaceutical
students, lectures that were published as “Grundtræk af forelesninger over system-
atisk botanik” (“Outlines of lectures in systematic botany”), in 1896.
What brought Warming back to Copenhagen and the inner circle of Danish
scholars was the desire of the King’s council to map and explore the natural
resources within the Danish Empire, such as those of Greenland, Faeroe Islands,
and eventually Iceland. These explorations started in the winter of 1884 with a
voyage to investigate the botany of Greenland. The sparse vegetation in this arctic
32923 Plant Community, Plantesamfund
landscape allowed Warming to achieve a swift and effective overview. The bare
landscape gave him an opportunity to understand the geographical distribution of
plants and see them in view of other plants and the entire habitat (Warming 1890).
The possibility of seeing plant communities in relation to the ecological environment
as a whole was an exciting turn in his research, and it became the methodology for
organizing his Brazilian material published as “Lagoa Santa” in 1892. Botanical
investigations into natural resources on contested Danish territory would occupy
much of Warming’s later work, such as in “Botany of the Færöes” (1901–1908)
and in “Botany of Iceland” (1912–1918). The aim of these investigations was to
establish Danish hegemony in the territories and open up for exploitation of plant
communities. This correlation between botany and resource management was
not accidental; Warming and his students sought to develop an ecological method
suitable for Danish social control in a foreign region (Christiansen 1924–1926,
pp. 799–800, 806–832).
Most of Warming’s work prior to the Greenland expedition was morphological
in content and systematic in outline. This research was widely respected among his
fellow scholars, but it did not attract students’ attention beyond preparing for
exams. What created excitement among the young was Warming’s introduction and
development of the plant community and other ecological concepts. Throughout the
1890s, the plant community became a particularly central concept. It was to explain
how plants could live and evolve together in “commensalism” without the dreadful
struggle for existence described by Charles Darwin (1809–1882). Warming’s
religious neo-Lamarckian views implied that plant communities were in stable
harmony slowly progressing towards higher development. This mirrored the con-
cept of community and progress in the Danish society.
The original Danish edition of “Plantesamfund” from 1895 was a short book
meant to provide the reader with a sense of overview. The book grew with each
translation, however, since Warming continuously added more details to substanti-
ate his claims. The key terms and concepts, however, hardly changed in the new
versions of the book. The German and English translations were thus conceptually
similar, while the number of examples and elaborations grew with each volume.
Most non-Danish scholars learned about the plant community concept through the
widely read German translation of “Plantesamfund” by Emil Knoblauch which was
published as “Lehrbuch der ökologischen Pflanzengeographie” in 1896. In Britain,
Tansley, for example, thought that Warming “opened [...] a new way of looking at
the plant world” (Tansley 1924, p. 54) and he adapted the German translation as a
textbook for his own botany classes at the University College, London. The Central
Committee for the Survey of British Vegetation (with Tansley as Chairman) would
dedicate their famous “Types of British Vegetation” from 1911 to Warming as “the
father of modern plant ecology.” It was the Edinburgh ecologist Balfour who
arranged for the first English translation. Warming wrote a fully revised manu-
script; he also upgraded the morphological content by leaning on the German ver-
sions. Balfour thus claimed in 1909 (with Warming as the authority and to Tansley’s
annoyance), that that University of Edinburgh was the center for plant community
and ecological research.
330 P. Anker
Henry C. Cowles (1869–1939) at University of Chicago was much pleased with
the morphological turn of ecological methodology, while Tansley wrote a long
critical review of the English edition, in which he advised his readers to stick to the
first German version (Cowles 1909; Tansley 1909). The tension among British
ecologists with regard to how to read Warming soon evolved into a major debate
between the mechanist inspired ecology of Tansley and the holistic “biotic
community” reading of Warming by Balfour’s student Phillips (Phillips 1931;
Anker 2001). It was the ordering of plants by Warming according to geographical
factors that intrigued Tansley. He was not against morphology as such when he first
read Warming. The tension that developed between him and Balfour first became
apparent when Tansley learned to appreciate genetic and biochemical research
around 1901. Tansley now started to promote genetics and plant geography as the
right ecological approach, while Balfour stuck to the morphological study of
tracing the ancestral history of species as a methodological basis of ecology.
Warming himself did not believe in the value of genetics, and the subsequent
German editions of his book grew with a 600 page morphological enlargement in
its final version of 1918. (Warming 1909, p. vi; Warming and Graebner 1918;
Goodland 1975). Tansley would later renew his interest in Danish botany by study-
ing the work of the Warming student Christen Raunkiaer (1860–1938), who over
the years had elaborated on Warming’s plant community concept in the direction of
functional classification in what he called “life-form-systems” of plants. Tansley
initiated a translation of Raunkiaer’s collected papers to English, which appeared
in 1934 (Raunkiaer 1934). Raunkiaer emphasized the importance of statistical
methods in studying plant communities, something that caught Tansley’s attention
while working out his own “ecosystem” concept (Tansley 1935).
References
Anker P (2001) Imperial ecology: environmental order in the British Empire, 1895-1945. Harvard
University Press, Cambridge
Christiansen C (1924) Den Danske botaniks historie. Hagerups Forlag, Kopenhagen
Coleman W (1986) Evolution into ecology? The strategy of Warming’s ecological plant geography.
J Hist Ecol 19:181–196
Cowles HC (1909) Ecology of plants. Bota Gaz 48:149–152, 465–466
Goodland RJ (1975) The tropical origin of ecology: Eugen Warming’s Jubilee. Oikos 26:240–245
McIntosh RP (1985) The background of ecology. Cambridge University Press, Cambridge
Nicolson M (1983) The development of plant ecology 1790–1960. Ph.D. thesis, University of
Edinburgh, Edinburgh
Phillips J (1931) The biotic community. J Ecol 19:1–24
Prytz S (1984) Warming: Botaniker og Rejsende. Bogan, Kopenhagen
Raunkiaer C (1934) The life forms of plants and statistical plant geography. Clarendon, Oxford
Rosenvinge LK, Warming E (eds) (1912–1932) The botany of Iceland, 4–5th edn. J. Frimodt,
Kopenhagen
Rosenvinge LK, Warming E (eds) (1912–1932). The botany of Iceland Vols. 4–5. J. Frimodt,
Kopenhagen
Schimper AFW (1898) Pflanzengeographie auf physiologischer Grundlage. Fischer, Jena
33123 Plant Community, Plantesamfund
Smuts JC (1926) Holism and evolution. Macmillan, London
Söderqvist T (1986) The ecologists: from Merry naturalists to Saviours of the Nation. Almquist &
Wiksell, Stockholm
Tansley AG (1909) Oecology of plants. New Phytol 8:218–227
Tansley AG (1911) Types of British vegetation. Cambridge University Press, Cambridge
Tansley AG (1924) Eug. Warming in memorian. Bot Tidsskr 39:45–56
Tansley AG (1935) The use and the abuse of vegetational concepts and terms. Ecology 16:284–307
Warming E (1879) Haandbok i den systematiske botanik. Philipsens Forlag, Kopenhagen
Warming E (1880) Den almindelige botanik. Philipsens Forlag, Kopenhagen
Warming E (1890) Botaniske exkursioner. Hovedbiblioteket, Kopenhagen
Warming E (1892) Lagoa Santa: Et bidrag til den biologiske Plantegeografi. Bianco Lunos Kgl.
Hof-Bogtrykkeri, Kopenhagen
Warming E (1895a) Plantesamfund: Grundtræk af den økologiske plantegeografi. Philipsens
Forlag, Kopenhagen
Warming E (1895b) A handbook of systematic botany. Swan Sonnenschin, London
Warming E (1896a) Lehrbuch der ökologischen Pflanzengeographie. Gebrüder Borntrager,
Berlin
Warming E (1896b) Grundtræk af forelesninger over systematisk botanik. Det Nordiske Forlag,
Kopenhagen
Warming E (1909) Oecology of plants: an introduction to the study of plant-communities. Oxford
University Press, London
Warming E, Graebner P (1918) Eug. Warming’s lehrbuch de ökologischen pflanzengeographie.
Gebrüder Borntraeger, Berlin
Warming E (1901–1908) Botany of færöes, 1st–3rd edn. Det Nordiske Forlag, Kopenhagen
333
Chapter 24
Looking at Russian Ecology Through
the Biosphere Theory
Georgy S. Levit
A. Schwarz and K. Jax (eds.), Ecology Revisited: Reflecting on Concepts, Advancing Science,
DOI 10.1007/978-90-481-9744-6_24, © Springer Science+Business Media B.V. 2011
Introduction
The biosphere theory is crucial for all environmental sciences including scientific
ecology. In Russia, the theory was from the very beginning a powerful factor affect-
ing global and other holistic approaches in the life sciences. The theory was
invented by Vladimir Ivanovich Vernadsky (1863–1945), who is regarded as one of
the most famous Russian naturalists. In the history of Russian science he is referred
to as a “savant” and influential thinker in rather different fields such as biogeo-
chemistry, radiogeology, or crystallography, and also philosophy of science. In
recent times Vernadsky is becoming appreciated also in the Western world. James
Lovelock, author of the Gaia-theory, wrote: “We discovered him to be our most
illustrious predecessor” (Lovelock 1986).
Therefore it is all the more astonishing that the origins of the theory and its
perception by different scientific communities, such as ecologists, biochemists or
geographers, are not very well known. This is even more true for the different
national, language and institutional contexts. At present there are about 1,000 pub-
lished works about Vernadsky, but his biosphere theory has not been adequately
investigated, reconstructed and appreciated. One reason might be the ideological
pressure and censorship in the former USSR. Another reason is the complexity and
quantity of his scientific literature: Vernadsky produced about 200 articles an books
in several languages directly connected to the biosphere theme and living matter.
Concerning the presence of Vernadsky in the history of science there is a rather
paradoxical situation: In the Russian history of science Vernadsky is a super star of
natural science and philosophy. His fame and influence certainly can be compared
with that of Ernst Haeckel in the German speaking countries. This contrasts sharply
to the position of Vernadsky in history of science in the West, where he is rather
underevaluated or even ignored (Ghilarov 1995), which is also true for the commu-
nity of Western biogeochemists that mostly do not know the founder of their own
G.S. Levit (*)
History of Science & Technology Program, University of King’s College, 6350 Coburg Rd.,
Halifax, NS, Canada, B3H 2A1
e-mail: Georgy.Levit@ukings.ns.ca
334 G.S. Levit
discipline. As the Vernadsky specialist Andrei Lapo claims: “To put the contempo-
rary situation concerning Vernadsky’s popularity on a world-wide scale in a graphic
phrase, one can say that the dinner table has already been laid, but the guests are
arriving late” (Lapo 2001). Yet Vernadsky’s heritage is of crucial interest not only for
geochemists, but also for ecologists, since “we should also credit Vladimir Vernadsky
with the title of father of the global ecology [...]” (Grinevald 1996, p. 48).
The following study seeks to point at some of these shortcomings. The first part
(Vernadsky and the Russian Science through the first half of the 20th century) gives
some insights into the disciplinary and institutional influence of Vernadsky and the
specific socio-political situation in Russia. In the second part (the essentials of the
biosphere theory) the biosphere theory itself will be presented. The last part
(Vernadsky’s impact on ecology and global sciences) gives some examples illustrat-
ing the reception of the biosphere theory in the Russian- and non-Russian-speaking
scientific communities.
Vernadsky and the Russian Science through the First Half
of the Twentieth Century
Initial empirical impulse for the biosphere theory was the idea of interconnected-
ness and lawfulness of geological processes, accompanied by the idea of interplay
between living and inert processes. The former is illustrated in Vernadsky’s early
works, e.g. by paragenesis, i.e. the regularities of mineral formation in an ore
deposit. Already in the very early beginnings of developing the theory, Vernadsky
founded a new scientific school detached from mineralogy and soil science. At that
same time the American scientist Frank Wigglesworth Clarke (1847–1931) pub-
lished comparable ideas in his “Data of Geochemistry” (Clarke 1908). However, in
contrast to Clarke, Vernadsky paid a lot attention to the role played by living matter
in the history of the earth’s crust and the atmosphere. Already in 1909 Vernadsky
delivered a paper to the “Meeting of the Russian Naturalists and Physicians” on the
basic principles of a new science geochemistry (Aksenov 1994, p. 111).
At the same time Vernadsky was beginning to work in the field of radioactiv-
ity. In 1908 he took part in a conference sponsored by the “British Association
for the Advancement of Science”1, where he met John Joly, one of the pioneers
of radioactivity research. Vernadsky was deeply impressed by the report of Joly
and already in 1909 founded the first radiological laboratory in Russia. The com-
bination of radioactivity studies with the idea of interconnectedness of living
organisms ultimately (after the WWII) led to the occurrence of Russian
radioecology.
1 Vernadsky was a member of this Association since 1889.
33524 Looking at Russian Ecology Through the Biosphere Theory
Some Biographical Notes
Vladimir Ivanovich Vernadsky (1863–1945) originates from the centre of Russian
cultural and academic life.2 In 1881 Vernadsky enrolled at St. Petersburg University,
where he was a student of such brilliant scientists as the chemists
A. Butlerov (1828–1886) and D. Mendeleev (1834–1907), the botanist A. Beketov
(1825–1902), the zoologist N. Wagner (1829–1907) and the physiologist
I. Sechenov (1829–1905). The greatest influence on Vernadsky had however the
soil scientist and mineralogist Vassilij Dokuchaev (1846–1903), the founder of
modern soil science and genetic pedology. Dokuchaev founded a landscape science
as a part of physical geography and created a concept of the natural climate related
zones. Moreover, it is recognised now that he was the first to declare the necessity
of a new synthetic science for studying “the genetic, eternal, lawful interconnec-
tions existing between the forces and bodies of inert and living nature” (Dokuchaev
1898). Thus Dukochaev can be also regarded as a fore-runner of the modern
ecosystem approach in the life sciences.
Vernadsky completed his examinations for the degree of candidate of science in
mineralogy and “geognosia”, and in 1888 left St. Petersburg for Germany. Germany
was regarded as the strongest academic land and German language was the lan-
guage of international scientific publications. Vernadsky decided to study crystal-
lography under the supervision of Paul Groth (1843–1927), who had a Chair in
Mineralogy at the University of Munich.
In 1889 Vernadsky moved from Munich to Paris where he started to work simul-
taneously under the guidance of the chemist Henry Le Chatelier (1850–1936) and
the mineralogist Ferdinand Fouqué (1828–1904). Le Chatelier helped Vernadsky to
find his dissertation topic and, as Vernadsky later recognised, significantly influ-
enced his scientific work. Working with the problem of crystalline polymorphism
Le Chatelier indirectly contributed to Vernadsky’s later space-time and biosphere
theories. In 1890 Vernadsky returned to Russia and after completing two disserta-
tions (master and Ph.D.) and in 1902 ultimately got a chair for mineralogy and
crystallography at the Moscow University.
In 1910 Vernadsky visited Eduard Suess (1831–1914) in Vienna. Suess was the
first scientist to use the term “biosphere” in the sense close to our modern usage,
however without proposing a clear concept underlying the term. Already in 1911
this term appeared in the work of Vernadsky although without definition. The
crucial step was made in 1912 as Vernadsky published a programmatic paper “On
Gaseous Exchange of the Earth’s Crust”, where he emphasized that almost all of
the earth’s gases are of biogenic nature and involved in cyclical processes
2 Vernadsky was born in the capital of the Russian Empire Sankt-Petersburg into the family of
Ivan Vernadsky (1821–1884), a professor of economics and statistics in the Alexandrovsky Lyce,
the elite college-like school, where, for example, the best known Russian poet Alexander Pushkin
was educated.
336 G.S. Levit
(Vernadsky 1912). 60 years later the same observation delivered empirical basis of
the so-called Gaia-hypothesis proposed by the English inventor James Lovelock
(e.g. Lovelock 1972; Lovelock and Margulis 1974; Levit 2000). This was also the
turning point in Vernadsky’s orientation to biological phenomena. But in contrast
to a purely biological approach, he began to think of life in terms of global geo-
chemistry. After the October (Bolshevik) revolution (1917) Vernadsky moved to
Kiev (nowadays the capital city of the Ukraine), where he was elected as the first
President of the Ukrainian Academy of Science. In the same year he initiated
biogeochemical scientific investigations. At the initial stages of this work
Vernadsky formulated the basic tasks of the newly established science (Lapo and
Smyslov 1989, p. 55): (1) to calculate a quantitative elementary composition of the
different species; (2) to investigate the geochemical history of silicon, copper,
zinc, lead, silver and some other elements; (3) to determine some other geochemi-
cal characteristics of living organisms like the average weight and water content
as well as the percentage of carbon in the organisms.
In 1921 Vernadsky received a letter from the Rector of the Sorbonne University
(Paris) with an invitation to teach a course on geochemistry and in 1922 Vernadsky
arrived in Paris where he made crucial steps in the direction of the biosphere theory.
Supported by the foundation of R. Rosenthal (a French “pears king” of Russian
origin) Vernadsky laid the baselines for his “The Biosphere” (1926). In 1923
Vernadsky for the first time used the very term “biogeochemistry” (Mochalov 1982,
p. 242).
Establishing and Widening the Concept in Russia
In March 1926 Vernadsky returned to Leningrad (St. Petersburg). Since he was well
known for his anti-communist views, his decision to come back to Soviet Russia
puzzled his biographers. Based on the archival materials of the Bakhmeteff Archive
at the Columbia University, Bailes (1990) and Kolchinsky and Kozulina (1998)
arrived at the conclusion that Vernadsky did make a considerable effort to remain in
the West. Vernadsky was in no way a sympathizer of the Soviet authorities. In the
time between two Russian revolutions (February and October 1917) he was involved
in the anti-Bolsheviki resistance as a member of the Constitutional Democrats Party.
However, he was unable to obtain permanent sufficient funding of his grandiose and
expensive biogeochemical research program in the Western countries. Vernadsky
decided to return to the USSR, realising that this was the only way to fullfill his
scientific mission (e.g. Vernadsky 1998). Paradoxically the totalitarian state in
Russia offered in this case more possibilities than institutions of the liberal world.
Although after the October Revolution 1917 the land was plunged into chaos and
economic disorder, it was at the same time the period of intensive innovations and
foundations of new research institutes in the life sciences. New authorities were
looking for a new scientifically based secular ideology. Life sciences and especially
Darwinism were seen as basic for the Marxist worldview (Kolchinsky 2006, p. 273).
33724 Looking at Russian Ecology Through the Biosphere Theory
So Nikolaj Koltzov (1872–1940) was able to found in Moscow an Institute for
Experimental Biology. In St. Petersburg Nikolaj Vavilov (1887–1943) initiated an
Institute for Plant Studies known nowadays as the Nikolai Vavilov Institute of Plant
Growing and Research. Another example is Alexei N. Sewertzoff (1866–1936), the
founder of Darwinian evolutionary morphology, who in 1922 established a
Department of Zoology at the University of Moscow, and seven years later (1930) a
Laboratory of Evolutionary Morphology (as a structural unit of the Institute of
Comparative Anatomy at Moscow University). In 1934, based on his Laboratory and
including the Institute of Palaeontology, Sewertzoff founded the Institute of
Evolutionary Morphology, which two years later was divided again into the Institute
of Palaeontology and the Institute of Evolutionary Morphology, which was later
reorganized into the A. N. Sewertzoff Institute of Ecology and Evolution. With more
then 700 members, the institute is now one of the biggest ecological institutes
worldwide.
Vernadsky initiated various institutions. For example, in 1926 he established a
Commission for the History of Knowledge (1926–1932) which later, after some
reorganisations, was transformed into the Institute of History of Natural Science
and Technology (1946), still in existence. Two years later (1928) the official foun-
dation of the Vernadsky’s Biogeochemical Laboratory of the Academy of Science
(BIOGEL) took place. After the series of reorganisations BIOGEL (1947) became
a part of the V.I. Vernadsky Institute of Geochemistry and Analytical Chemistry
with an emphasis on ecological-biogeochemical research, such as the ecological
assessment of biogeocenoses. At the same time Vernadsky conducted his scientific
research and in 1926 presented his views on the biosphere in the book of the same
name (“Biosfera”) published in Russian in Leningrad. Three years later the book
was translated and published in French (Vernadsky 1929).
All these scientific developments took place in the cotext of political repressions
and strengthening totalitarianism in the Soviet Russia. In 1928, with the end of the
liberal economic policy (so-called New Economic Policy) and the emergence of
forced of forced collectivization and industrialisation known as Stalin’s Great
Break, the repressive machine came into the a much more intensive phase.
It is astonishing that in spite of this accelerating terror machine Vernadsky
enjoyed the liberty of traveling abroad. Thus Vernadsky spent the summer of 1929
in Germany and Czechoslovakia. In 1932–1933 he travelled in various countries
including Germany, France, the UK, Poland and Czechoslovakia. In Münster he
contributed a paper (1932) “Die Radioaktivität und die neuen Probleme der
Geologie” (Radioactivity and new problems of geology) to the “Deutsche Bunsen-
Gesellschaft für Physikalische Chemie” (German Bunsen-Society for Physical
Chemistry). In England, Vernadsky communicated with Frederick Soddy (1877–
1956) who founded a theory of isotopes. The study of the isotopic composition and
radioactive elements in living matter became since that moment an important line
of Vernadsky’s research. In the 1930s Vernadsky also continued publishing his
papers abroad (for example: Vernadsky 1930, 1934, 1935). It must be emphasized
here that Vernadsky was not an absolute exception from the rule. As already men-
tioned, Sewertzoff published his major evolutionary book first in Germany
338 G.S. Levit
(Sewertzoff 1931) and only then in Russian, in the Soviet Union. The isolation of
the soviet scientists in that period was not of an absolute nature. Table 1 presents,
the publications of the Soviet scientists in Germany in only one, although central
German biological publication. The table shows that even after the totalitarian
regimes were established both in USSR and Germany there was no impenetrable
borderline between the scientists of two countries. The publications of the Soviet
scientists cease only after the beginning of the WWII.
In February, 1934 Sergej Oldenburg (1863–1934), the Permanent Secretary of
the Academy of Science, died. He had been Vernadsky’s closest friend and a strong
supporter. His death symbolized also the end of the Petersburg period of the
Academy of Science. Vernadsky’s Biogeochemical Laboratory moved to Moscow
together with the Academy. Approximately at that time Vernadsky came to the idea
of writing a book where his holistic views on nature would be expressed both scien-
tifically and philosophically. By 1936 Vernadsky understood that this was impossi-
ble to do in one work, and decided to separate this task into two books, one
philosophical and one strictly scientific. Thus Vernadsky began to work on his main
works “The Chemical Structure of the Earth’s Biosphere and Its Environment”,
which was published only 20 years after his death, and laid foundations for modern
biogeochemistry and global ecology (Vernadsky 1965). After the war with Germany
broke out (22 June, 1941) Vernadsky was evacuated to the health resort Borovoje in
Kazakhstan. The two years in Borovoje were highly productive. Vernadsky wrote
the important third issue of series “Problems of Biogeochemistry” (1980) which he
saw as his scientific will. He worked also on his main generalizing work “The
Chemical Structure of the Earth’s Biosphere and Its Environment” where his basic
claims, expressed first in “The Biosphere” were revised and developed.
In Borovoje many outstanding scientists thus came to live and work together such
as, for example, the founder of the nomogenesis theory (theory of directed evolu-
tion) and the theory of geographic zones Leo (Lew) S. Berg (1876–1950) (Levit and
Hoßfeld 2005), the founder of the theory of biogeocenosis as an elementary unit of
the biosphere, Vladimir Sukachev (1880–1967), and one of the architects of Russian
evolutionary synthesis Ivan Schmalhausen (1884–1963). As is clearly documented
they all experienced Vernadsky’s influence and all of them performed a crucial role
in the growth of Russian life sciences, including various branches of ecology.
The Essentials of Vernadsky’s Biosphere Theory
In the theoretical system of Vernadsky, the concept of the biosphere is required by the
new branch of science created by himself: biogeochemistry. Biogeochemistry studies
the geological manifestations of life and considers biochemical processes in living
organisms in relation to their impact on the geosphere (Vernadsky 1997, p. 156).
The competence of biogeochemistry is defined, on the one hand, by the geological
manifestations of life taking place under this aspect, and on the other, by the internal
biochemical processes in the organisms the living population of our planet. In both cases
33924 Looking at Russian Ecology Through the Biosphere Theory
(for biogeochemistry is a part of geochemistry) one may identify as study objects not only
chemical elements, i.e. the usual mixtures of isotopes, but also various isotopes of one and
the same chemical element.3
Thus the specificity of biogeochemistry, in relation to classic geochemistry,
includes its concentration on living matter as the major factor in biogenic migration
of chemical elements. Neither living organisms by themselves nor their environ-
ment abstracted from them are, Vernadsky argued, the specific objects of biogeo-
chemistry. A biogeochemist is interested, first of all, in studying the cyclic processes
of the exchange of chemical elements between living organisms and their environ-
ment. The latter can only be described on the basis of a detailed study of interrela-
tions of living and inert (non-living) matter in the space-time of Earth and
throughout the history of Earth. How can the main subject of biogeochemical
research be defined? Biogeochemistry never aims at the organismic level or at the
environmental level alone. It concentrates, in Vernadsky’s words, on the biologi-
cally controlled flow of atoms, which takes place in a specific geological domain.
In order to define this specific geological domain as the research field of the
newly created science, biogeochemistry, Vernadsky introduced his interpretation of
the term biosphere. The biosphere of the Earth appears as a geosphere occupied and
organised by life and thus can be seen as a geological envelope.
Being a geological envelope, the biosphere can be also structured geologically
(Vernadsky 1991, p. 120):
The biosphere appears in biogeochemistry as a peculiar envelope of the Earth clearly dis-
tinct from the other envelopes of our planet. The biosphere consists of some concentric
contiguous formations surrounding the whole Earth and called geospheres. The biosphere
has possessed this perfectly definite structure for billions of years. This structure is tied up
with the active participation of life, is conditioned by life to a significant degree and is
primarily characterised by dynamically mobile, stable, geologically durable equilibria
which, in distinction to the mechanical structures are quantitatively fluctuating within
certain limits in relation to both space and time.
Under the various “geospheres” Vernadsky (1965, pp. 107–108) understands the
troposphere, the hydrosphere, the land surface and the sphere of the subterranean
life. However, Vernadsky’s approach to the biosphere goes far beyond the purely
stratigraphical statements. Examining living matter from the biogeochemical view-
point, Vernadsky (1994b) arrived at the conclusion that the chemical compounds of
the different species do not reflect that of their environment, but, on the contrary,
living matter has determined the geochemical history of almost all the elements of
the Earth’s crust in the process of making the environment favourable to itself.
Thus, living matter shapes the biosphere into a self-regulating system. The bio-
sphere, being seen as a self-regulating system, embraces both the totality of living
organisms (living matter) and their environment to the extent it is involved in the
actual processes of life, that is, including the troposphere, the ocean, and the upper
3 Vernadsky began the work on the “Scientific Thought as a planetary Phenomenon” (the book
I quote here) in the late 1930s. The book was published in an uncensored form only in 1991.
In 1997 appeared the English translation.
340 G.S. Levit
envelopes of the Earth crust, possibly down to the mantle. The structure of the
biosphere is described as a dynamic equilibrium: “Not a single point of this system
is fixed during the course of geological time. All points oscillate around a certain
midpoint” (Vernadsky 1997, pp. 225–227).
A good example of such dynamic equilibrium is the troposphere. Vernadsky
claimed that “all basic gases of the troposphere and of the higher gaseous envelopes
N2, O2, CO2, H2S, CH4, etc., are produced and quantitatively balanced by the total
activity of living matter. Their sum total is quantitatively invariable over geological
time [...]” (1965, p. 238). Thus, Vernadsky concludes, “life, i.e. living matter creates
the troposphere and constantly maintains it in a specific dynamic equilibrium.” It
can be remarked here, that the first Gaian principle of atmospheric regulation
(Lovelock and Margulis 1974) actually was derived by Vernadsky on the basis of his
biogeochemical research 50 years before Lovelock (Levit and Krumbein 2000).
However the biosphere is, in Vernadsky’s view, not only a self-regulating, but
also an evolving system: “We can and must talk about the evolutionary process of
the biosphere by itself” (Vernadsky 1991, p. 20).
Based on his experimental work, Vernadsky already in the beginning of the
1920s concluded that his concept of living matter would influence the evolutionary
theory. Studying the natural history of the chemical elements, Vernadsky (1994a,
pp. 66–68) arrived at the conclusion that living matter modifies the environment.
Living matter, in its turn, is determined by the inert environments.
An important characteristic of the biosphere is its holistic nature, which is guaran-
tied by the various biogeochemical functions. A biogeochemical function is a role
which a taxon performs in the biospheric cycles. The major biogeochemical functions
include, according to Vernadsky, five groups: (1) The gas-functions, regulating the
gaseous structure of the atmosphere including submarine and subterranean environ-
ments. (2) The function of concentration, i.e. the ability of organisms to capture and
concentrate the chemical elements of their environments. Concentration functions of
the first kind describe accumulation by the organisms of elements composing all living
organisms without exception; the functions of the second kind can be fulfilled only by
the certain kinds of organisms, playing thus an unique role in the trophic chains, as for
example molluscs concentrating heavy metals. (3) The oxidation-reduction functions;
(4) the biochemical functions of organisms generating biogenic migrations of atoms
connected with feeding, breathing, multiplication and destruction of organisms; (5)
the biogeochemical functions of the mankind (Vernadsky 1965, p. 237).
Vernadsky has also shown how exactly different biogeochemical functions cor-
respond to specific taxonomic groups. For example, the oxidising function is car-
ried out by autotrophic bacteria and the function of destruction of organic
compounds by chemoorganotrophic bacteria and fungi. Analysing these results
Vernadsky came to the following three conclusions:
All basic biogeochemical functions can be carried out by unicellular organisms; –
There is no species able carry out all these functions; –
In the course of geological time, different species may have replaced one –
another, but the biogeochemical functions must have been carried out.
34124 Looking at Russian Ecology Through the Biosphere Theory
Ultimately this means that life must have been occurred in the form of a
biosphere-like from the very beginning and can exist only in the form of the biosphere,
since various biogeochemical functions have to be fulfilled simultaneously: “The
first occurrence of life in the biosphere could not be in form of separate organisms
but only in the form of the sum total of organisms carrying out various geochemical
functions. Biocoenoses necessarily had to occur from the very beginning”
(Vernadsky 1994b, p. 459).
On these grounds Vernadsky negated the polyphyletic evolutionary model for the
very first stages of biospheric evolution. In his late works, Vernadsky claimed that the
biosphere as a self-regulating system has its clearly definable evolutionary “interests”.
A leading force of the evolution of the biosphere is living matter, which has its own
process of evolution partially independent from the needs of adaptation. The bio-
sphere as a whole behaves itself as if it had a peculiar evolutionary strategy: “We can
and must talk about the evolutionary process of the biosphere by itself” (Vernadsky
1991, p. 20; 1997, p. 30). One of the basic methods of realisation of these “interests”
is increasing the intensity and complexity of the biogenic migration of atoms.
Vernadsky’s Impact on Ecology and Global Sciences
Vernadsky created a theoretical system that influenced the whole range of environ-
mental sciences. The most evident is Vernadsky’s influence in geochemistry and
biogeochemistry. Vernadsky understood geochemistry as a natural history of ter-
restrial chemical elements. It is however important that his approach allowed scien-
tific predictions about the ways of migration of the chemical elements including
their compatibility in the various kinds of rocks. This was significant also for the
applied geology, e.g. for the minerals search. One of the most known followers of
Vernadsky in the field of geochemistry was his student Alexander Fersman (1883–
1945), who gave the first regular course of general geochemistry as early as 1911
and became a founder of an influential scientific school. After Vernadsky’s and
Fersman’s death the Russian school of geochemistry was led by Alexandr
Vinogradov (1895–1975). Vinogradov was Vernadsky’s deputy in the biogeochemi-
cal laboratory and after the WWII organised and headed the Vernadsky Institute
of Geochemistry and Analytical Chemistry (1947–1975). He is also regarded as
the major follower of Vernadsky in the field of biogeochemistry. In contrast to
geochemists concentrating on migration of chemical elements and their composi-
tion in the rocks and minerals, biogeochemistry involves the study of cyclic bio-
genic migrations of chemical elements caused by activity of living organisms
(Vinogradov 1953). Following Vernadsky Vinogradov proposed that the chemical
composition of living organisms is a result of biological evolution, which proceeded
in certain environments.
Along similar lines, based on Vernadsky’s and Vinogradov’s concept of biogeo-
chemical provinces (see above), Victor Kovalsky (1899–1984) coined a concept of
geochemical ecology (Kovalsky 1974). Kovalsky proceeded from the assumption
342 G.S. Levit
that the biosphere is a biogeochemically heterogeneous body and that the chemical
composition of elements is part of their adaptation to the environment. He showed
that different species develop different strategies adapting to the specific environ-
ments. For example, Pyrethrum parthenifolium in the molybdenum reach soils of
Armenia adapts by reducing molybdenum accumulation, while leguminous plants,
on the contrary, increase molybdenum concentration.
An important line of Vernadsky’s influence was his impact on the co-architect
of the Modern Synthesis, the German-Russian biologist Nikolai Timofeév-
Ressovsky (1900–1981).4 In his own words, he was, first of all, interested in
Vernadsky’s biosphere theory. Supporting the idea of biogeocenoses as structural
units of the biosphere, he wrote: “The biosphere in its entirety consists of more or
less complex biotic and abiotic components, i.e. biogeocenoses. In other words, the
biogeocenoses are the precise environments in which the evolutionary process of
any group of living organisms takes place” (Timoféev-Ressovsky et al. 1975,
p. 249). Timoféev-Ressovsky sought to create a new branch of evolutionary theory
studying evolution of biogeocenoses (ecosytsems) and the biosphere. Specifically,
following his forceful repatriation after WWII, Timoféev-Ressovsky founded a
school of radiation biogeocenology and radiation biogeochemistry (Tjurjukanov
and Fiodorov 1996, pp. 97–98). He defined biogeocenoses as “dynamic systems,
which at the same time can be in a state of dynamic equilibrium over quite a long
biological time period (in the course of many generations of living beings residing
in this beogeocenosis)” (Timoféev-Ressovsky et al. 1975, p. 309). The biosphere is
defined as the sum total of biogeocenoses. Timoféev-Ressovsky insisted that there
is a significant difference to the term ecosystem, predominantly used in the Western
world, because biogeocenosis comprises all abiotic factors and all biotic dependen-
cies in a relatively isolated system occupying clearly detectable zones (e.g. a pine
forest or a swamp). To study the input, output and cyclicity of these systems
Timoféev-Ressovsky proposed using nuclear markers. Also, he pioneered the
investigations into the impact of radioactivity on the living organisms and ecosys-
tems and thus initiated studies in radiation ecology.
The influence of Vernadsky’s biosphere theory and Timoféev-Ressovsky’s
vision of a new science studying evolution of the whole biogeocenoses influenced
also the very recent studies in evolution of multi-species communities and paleo-
ecology, as seen in the works of Vladimir V. Zherikhin (1945–2001), one of the
central figures in the field. Methodologically Zherikhin proceeded from the in
Russia unpopular organismic approach based on the ideas of Frederic E. Clements
(1874–1945), and developed further by Stanislav Razumovsky (1929–1983) (Rautian
2003). Zherikhin claimed that biological communities can be analysed from the
viewpoint of their “ontogeny” (endoecogenesis) and “phylogeny” (phyloecogensis).
As a result Zherikhin – with co-authors – proposed a “biotocenogenetic” model of
4 Timofeév-Ressovsky – who coined a somewhat awkward term “vernadskology” – wasn’t a direct
pupil of Vernadsky, although he met him two times in Berlin in the mid of 1920s (Timofeév-
Ressovsky 1995).
34324 Looking at Russian Ecology Through the Biosphere Theory
feedback loops between taxa and biocoenoses. The direction of phylogenesis in this
model will be, to an extent, directed by a biocenosis (biogeocenosis) so far the
included taxa evolve co-adapting by specialization and thus their further specialisa-
tion is then “predestined” by the whole system. Vernadsky is important here, first
of all, because of two points. First, the biosphere is the ultimate biogeocenosis and
a self-regulating system directing evolution of the lower systems. Second, Zherikhin
adapted Vernadsky’s thesis on living matter as the major mover of geochemical
cycles (Zherikhin 2003, p. 348).
The biosphere studies in Vernadsky’s sense were never interrupted in the
Russian speaking countries. In the most articulated contemporary version, repre-
sented – in my view – by Georgii Zavarzin (1997, 2003a, b), the biosphere theory
claims that phylogenetically independent prokaryotes are basic for the running of
biogeochemical cycles of the biosphere. This implies that (1) the Vernadskian
approach excludes strict monophily at the very early stages of biospheric evolution,
because life on Earth can exist only in the from of communities able to support
closed biogeochemical cycles (there were functionally complete microbial com-
munities even in the early Proterozoic); (2) evolution has an additive character
(“new” plus “old” and not “new” instead of “old”); (3) the Biosphere functions as
a well-balanced system of functionally complementary organisms, and the
Darwinian laws work only on the lowest level of this system.
Although the above examples can be in no way seen as exhausting the subject,
one can see that Vernadsky influenced nearly the whole range of global sciences in
Russia.
However, his influence is of a disproportional nature. In Russia Vernadsky
belongs to the pantheon of the most known and esteemed scientists. His name is to
be found even in school textbooks. His biosphere-noosphere theory is included in
the course “Basics of Natural Sciences” obligatory for all university students of all
disciplines. Not surprisingly his theory is studied in depth by students of philoso-
phy, evolutionary theory, ecology and, of course biogeochemistry. In the introduc-
tory chapter to a modern Russian university biogeochemistry textbook we read:
“Theoretical foundations of biogeochemistry are composed of the theories of living
matter and biosphere created by V.I. Vernadsky” (Dobrovolsky 1998)Vernadsky’s
ideas are also reflected in the very recent Russian governmental programs. One of
the explicit mentions of Vernadsky’s theories in the official documentation is found
in the 1996 Presidential decree concerning the “Concept of Russia’s Transition to
Sustainable Development” (Oldfield and Schaw 2006).
By contrast, Vernadsky remains little known in the Western world. There are few
contemporary researchers explicitly advocating Vernadsky’s views. The English
scientist James Lovelock and the American microbiologist Lynn Margulis (e.g.
Lovelock and Margulis 1974; Lovelock 1986, 1996; Margulis 1996) number
Vernadsky among their scientific “predecessors”, but repeatedly stress that they
came to their version of the biosphere theory (Gaia-hypothesis) independently of
Vernadsky. In addition, the only book available for these English-speaking scien-
tists is the recent translation of Vernadsky’s early work “The Biosphere” (Vernadsky
1998). Vernadsky’s major book “The chemical Structure of the Biosphere” remains
344 G.S. Levit
untranslated in Western European languages. Vernadsky’s follower, who clearly
declared Vernadsky’s theory as the basis of modern geophysiology, is the German
geomicrobiologist Wolfgang E. Krumbein (Krumbein and Schellnhuber 1992;
Krumbein and Lapo 1996). Vernadsky is mentioned in the recent introduction into
geomicrobiology (Ehrlich 2002).
Yet the majority of introductory English-language books on biogeochemistry
and global ecology do not even mention Vernadsky (Degens 1989; Schlesinger
1991, 2004; Libes 1992; Fenchel et al. 2000). The authors either somehow escape
the question on the origins of the biosphere concept and biogeochemistry, or begin
the story with the postwar developments. As Schlesinger in his most recent survey
paper on the “Global Change Ecology” puts it: “I mark the beginning of global
change science with the publication of ‘The Biosphere’ as a special issue of
Scientific American in 1970” (Schlesinger 2006). To fully realize the absurdity of
this situation one should imagine a book on evolutionary biology, which would
begin with Ernst Mayr and William Provine’s “Evolutionary Synthesis” (1980),
with no references to Darwin or with a few cursory remarks about him. At the same
time Vernadsky’s ideas penetrated the theoretical landscape without being actually
associated with his name. As the ecologist George E. Hutchinson (1903–1991) in
the above mentioned book claimed: “It is essentially Vernadsky’s concept of the
biosphere, developed about 50 years after Suess wrote, that we accept today”
(Hutchinson 1970). Yet in the situation of only fragmentary infiltration of
Vernadsky’s original concepts into the Western theoretical landscape, the proper
estimation of his theory by the international scientific community seems to be a
difficult task.
Summary
Vernadsky’s most important contribution to modern science was his grandiose
theory of the biosphere and living matter. The idea was developed through the first
half of the twentieth century and influenced the whole range of natural sciences
including the new field of ecology and evolutionary theory. Vernadsky understood
geochemistry as a natural history of terrestrial chemical elements. The specificity
of biogeochemistry, in relation to classic geochemistry, reflected his novel idea that
living matter was the major factor in the migration of chemical elements in the
biosphere. In Vernadsky’s terms, the biosphere is both a geological stratum and a
self-regulating system including both living organisms and their inert environ-
ments. Complemented by the concept of biogeocenosis coined (1940) by Vladimir
Sukachev, the biosphere appeared to be a self-regulating system consisting of bio-
geocenoses as its elementary structural units, which in their turn represent self-
regulating systems. Biogeocenoses comprise all abiotic factors and all biotic
dependencies in a relatively isolated natural system occupying a clearly detectable
zone, e.g., a pine forest or a swamp. This approach allowed scientific predictions
about the ways of migration of the chemical elements in the biosphere and became
34524 Looking at Russian Ecology Through the Biosphere Theory
crucially important for Russian ecology. A number of Russian scientists adopted
and extended Vernadsky’s new approach. These are, for instance, the concept of
geochemical ecology (Victor Kovalsky), the whole school of radiation ecology and
radiation biogeochemistry founded by Nikolai Timofeév-Ressovsky, important
impacts on modern paleoecology (also Sukachev and Timoféev-Ressovsky’s) and,
more recently, the approaches of Vladimir V. Zherikhin, the founder of the so-
called naturalistic microbiology. Alltogether, Vernadsky’s theoretical system was
one of the most crucial steps in shaping modern global sciences including
ecology.
Acknowledgement My research on the history of life sciences was supported by the Deutsche
Forschungsgemeinschaft (DFG) (Ho 2143/5–2). I am thankful to Astrid Schwarz and Christian
Haak for helpful comments on an earlier version of this paper.
References
Aksenov G (1994) On the scientific solitude of Vernadsky. Probl Philos 6:74–87 [in Russian]
Bailes KE (1990) Science and Russian culture in an age of revolutions: V.I. Vernadsky and his
scientific school, 1863–1945. Indiana University Press, Bloomigton/Indianapolis
Barrow J, Tipler F (1986) The anthropic cosmoplogical principle. Claderon Press, Oxford
Clarke FW (1908) Data of geochemistry. Government Printing Office, Washington, DC
Dana JD (1852) Crustacea, Reprinted in 1972. Antiquariat Junk, Lochem
Degens ET (1989) Perspectives on biogeochemistry. Springer, Berlin
Dobrovolsky VV (1998) Basics of biogeochemistry. Vyschaja Schkola, Moscow [in Russian]
Dokuchaev VV (1898) The concept of zones in nature, 2nd edn., 1948. Moscow Geografgiz, [in Russ]
Ehrlich HL (2002) Geomicrobiology, 4th edn. Marcel Dekker, New York
Fenchel T, King GM, Blackburn TH (2000) Bacterial biogeochemistry: the ecophysiology of
mineral cycling, 2nd edn. Academic, San Diego [u.a.], Reprinted
Fersman AE (1923) Khimitcheskije elementy zemli i kosmosa (Chemical Elements of the Earth
and the Cosmos). Khimtekhizdat, Petrograd
Ghilarov AM (1995) Vernadsky’s biosphere concept: An historical perspective. Q Rev Biol
70(2):193–203
Grinevald J (1996) Sketch for the History of the Idea of the Biosphere. In: Bunyard P (ed) Gaia
in Action. Floris Books, Edinburgh, pp 115–135
Hutchinson GE (1970) The biosphere. Sci Am 223(3):45–53
Kolchinsky EI (1990) The evolution of the biosphere. Nauka, Leningrad [in Russ]
Kolchinsky E, Kozulina A (1998) The burden of choice: why did V.I. Vernadsky return to the
Soviet Russia? Voprosy istorii estestvoznanija i tekhniki 3:3–25 [in Russian]
Kolchinsky EI (2006) Biology in Germany and Russia-USSR. Nestor-Istorija, St.-Petersburg [in
Russ]
Kovalsky VV (1974) Geokhimitcheskaja ekologija [Geochemical ecology]. Nauka, Moscow
Krumbein WE, Schellnhuber H-J (1992) Geophysiology of mineral deposits a model for a biologi-
cal driving force of global changes through Earth history. Terra Nova 4:351–362
Krumbein WE, Lapo A (1996) Vernadsky’s biosphere as a basis of geophysiology. In: Bunyard P
(ed) Gaia in action. Floris Books, Edinburgh, pp 115–135
Lapo AV, Smyslov AA (1989) Biogeochemistry: the foundations laid by V.I. Vernadsky. In:
Yanschin AL (ed) Scientific and social significance of Vernadsky’s creativity. Nauka, Moscow,
pp 54–61 [in Russian]
346 G.S. Levit
Lapo AV (2001) V.I. Vernadsky (1863–1945), the founder of the biosphere concept. Int Microbiol
4:47–49
Levit GS, Krumbein WE (2000) The biosphere theory of V.I. Vernadsky and the Gaia theory of J.
Lovelock: a comparative analysis of the two theories and two traditions. Zhurnal Obshchei
Biologii (J Gen Biol) 61(2):133–144
Levit GS (2001) Biogeochemistry, biosphere, noosphere: the growth of the theoretical system of
Vladimir Ivanovich Vernadsky (1863–1945), Series: “Studien zur Theorie der Biologie”
(Edited by Olaf Breidbach & Michael Weingarten). VWB-Verlag, Berlin
Levit GS, Hoßfeld U (2005) Die Nomogenese: Eine Evolutionstheorie jenseits des Darwinismus
und Lamarckismus. Verhandlungen zur Geschichte und Theorie der Biologie 11:367–388
Libes SM (1992) An introduction to marine biogeochemistry. Wiley, New York
Lovelock J (1972) Gaia as seen through the Atmosphere. Atmos Envir 6:579f
Lovelock J (1986) The biosphere. New Sci 1517:51
Lovelock J, Margulis L (1974) Atmospheric Homeostasis by and for the biosphere: the Gaia
hypothesis. Tellus 26:2–10
Margulis L, Sagan D (1995) What is life? A Peter N. Nevraumont Book, New York
Margulis L(1996) James Lovelock’s Gaia. In: P. Bunyard (ed) Gaia in action. Floris Books,
Edinburgh, pp 54–65
Mochalov II (1982) Vladimir Ivanovich Vernadsky. Nauka, Moscow
Oldfield JD, Schaw DJB (2006) V.I. Vernadsky and the noosphere concept: Russian understand-
ings of society-nature interaction. Geoforum 37(1):145–154
Por FD (1980) An ecological theory of animal progress – a revival of the philosophical role of
zoology. Perspect Biol 23(3):389–399
Rautian AS (2003) O nachalakh teorii evoliutzii mnogovidovykh soobstchestv i ee avtore (On the
beginnings of the theory of multi-species communities evolution – phylocenogenesis – and its
autor). In: Lubarsky G (ed) Zherikhin V.V. Izbrannyje trudy. KMK Press, Moscow, pp 1–42
Schlesinger WH (1991) Biogeochemistry: an analysis of global change. Academic, San Diego [u.a.]
Schlesinger WH (ed) (2004) Treatise on geochemistry – Vol. 8: Biogeochemistry. Elsevier
Pergamon, Amsterdam (u.a.)
Schlesinger WH (2006) Global change ecology. TREE 21(6):348–351
Sewertzoff A.N. (1931) Morphologische Gesetzmäßigkeiten der Evolution. Gustav Fischer
Verlag: Jena
Sytnik K, Apanovich E, Stoiko S (1988) V.I. Vernadsky. Life and activity in the Ukraine. Naukova
Dumka, Kiev [in Russian]
Teilhard de Chardin P (1961) The phenomenon of man. Harper & Row, New York/Evanston
Timofeév-Ressovsky NV (1995) Vospominanija (memoirs). Progress, Moscow
Timoféev-Ressovsky NW, Vononcov NN, Jablokov AN (1975) Kurzer Grundriss der
Evolutionstheorie. Gustav Fischer Verlag, Jena
Tjurjukanov AN, Fiodorov VM (1996) N.V. Timoféev-Ressovsky: Biosfernyje razdumja. AEN,
Moscow
Vernadsky VI (1902) O nauchnom mirovozzrenii (On the scientific worldview). Vorposy filosofii
i psikhologii 1(65):1409–1465
Vernadsky VI (1903) Osnovy kristallografii (The Fundamentals of Crystallography). Izdatelstvo
Moskovskogo Universiteta, Moscow
Vernadsky VI (1912) O gazovom obmene zemnoj kory (On gaseous exchange of the earth’s crust).
Izvestija Imp Akad Nauk Serija 6 6(2):141–162
Vernadsky VI (1924) La Géochemie. Alcan, Paris
Vernadsky VI (1926) Biosfera. NHTI, Leningrad
Vernadsky VI (1929) La Biosphère. Alcan: Paris
Vernadsky VI (1930) Geochemie in Ausgewählten Kapiteln. Autorisierte Übersetzung aus dem
Russischen von Dr. E. Kordes. Akademische Verlagsgesellschaft, Leipzig
Vernadsky VI (1934) Le Problème du Temps dans la Science Contemporaine. Revue Génerale des
Sciences Pures et Appliquees 45(20):550–558
Vernadsky VI (1935) Le Problème du Temps dans la Science Contemporaine. Revue Génerale des
Sciences Pures et Appliquees 46(7):208–213, 47(10): 308–312
34724 Looking at Russian Ecology Through the Biosphere Theory
Vernadsky VI (1944) Problems of biogeochemistry. (Trans: George Vernadsky Ed and condensed:
G E Hutchinson) Connecticut Academy of Arts and Sciences, New Haven [u.a.]
Vernadsky VI (1965) The chemical structure of the biosphere of the earth and of its environment.
Nauka, Moscow [in Russian]
Vernadsky VI (1980) Problems of biogeochemistry. III, BIOGEL. Nauka, Moscow [in Russian]
Vernadsky VI (1988) Philosophical thoughts of naturalist. Nauka, Moscow, p 520 [in Russian]
Vernadsky VI (1991) Scientific thought as a planetary phenomenon. Nauka, Moscow [in
Russian]
Vernadsky VI (1994a) Works on geochemistry. Nauka, Moscow [in Russian]
Vernadsky VI (1994b) Living matter and the biosphere. Nauka, Moscow [in Russian]
Vernadsky VI (1997) Scientific thought as a planetary phenomenon. Nongovernmental Ecological
V.I. Vernadsky Foundation, Moscow
Vernadsky VI (1998) The biosphere. A Peter A. Nevraumont Book, New York
Vinogradov AP (1953) The elementary chemical compositions of marine organisms. Memoir
Sears Foundation for Marine Research II. Yale University Press, New Haven
Vinogradov AP (1993) The geochemistry of isotopes and the problems of biogeochemistry:
selected works. Nauka, Moscow [in Russian]
Zavarzin GA (1997) The rise of the biosphere. Microbiology/Microbiology 6(66):603–611
Zavarzin GA (2003a) Evolution of the geosphere-biosphere system. Priroda 1:27–35 [in
Russian]
Zavarzin GA (2003b) Prirodovedcheskaja mikrobiologija (Naturalistic microbiology). Nauka,
Moscow
Zherikhin VV (2003) Izbrannyje trudy (selected papers). KMK Press, Moscow [in Russian]
Part VII
Border Zones of Scientific Ecology
and Other Fields
351
Introduction: The Core Paradigm of Geography
The major theme and the core theory of classical geography, that is, of mainstream
geography from the eighteenth to the twentieth century, can be summed up roughly as
follows: the interaction and symbiosis between regional modes of life and entire cul-
tures on the one hand and their concrete ecological milieu on the other. One could also
express it using some rather misleading set phrases, namely, the man-nature, man-space
or man-environment theme.1 This concrete ecological or physico-biotic milieu could be
taken to mean both the original milieu as well as that which had already been altered
through history. This was the “Rittersche Wissenschaft” which, from the time of Carl
Ritter (1779–1859) onwards and with regular and repeated reference to him, was to
bring together a broad range of “raw material” and to refine that material until it had
become a university subject suitable for research and teaching. Ritter’s “Erdkunde” has
had a major influence on many nations’ geographies, not least thanks to his numerous
students (and his students’ students) scattered throughout the world.
Ritter and his followers, however, were merely formulating a programme that
had already been an integral part of the common sense shared by educated people
in the 18th and early 19th century. From now on, this – to put it in modern terms –
“Kulturökologie” (cultural ecology) was no longer merely an almost ubiquitous
theme in the “educated common sense” and popular science of the time; it also
became the paradigm of a university discipline. As modern ecologists themselves
are aware, such a constellation does not bring undiluted benefits.
In the English-language literature, this man-and-environment theme in geography
is often simply called “the ecological tradition”.2 It is true that more recent
Chapter 25
Geography as Ecology
Gerhard Hard
1 On the theoretical concept of “Theoriekern” (theoretical core), synonymous with “Kerntheorie”
(core theory) and “Kernparadigma” (core paradigm) etc., cf. Stegmüller 1973, 1979, 1980.
2 Cf. e.g. Haggett 1965 (pp. 10 ff.) in the German edition of the book 1972 (p. 15) the term was
translated literally using “ökologisch” and “Ökologie” and was explained in typically allencom-
passing terms as follows: “Geographie als die Erforschung der Beziehungen zwischen der Erde
und dem Menschen” (Geography as the study of relationships between the earth and man).
G. Hard (*)
Institut für Geographie, Universität Osnabrück, Seminarstraße 19 a/b, D- 49069 Osnabrück, Germany
A. Schwarz and K. Jax (eds.), Ecology Revisited: Reflecting on Concepts, Advancing Science,
DOI 10.1007/978-90-481-9744-6_25, © Springer Science+Business Media B.V. 2011
352 G. Hard
English-language geography in particular contains more references to several
(at least four or five) different “traditions”, “schools”, “major themes” or “focal
points of interest” in geography’s past and present, with the aforementioned “eco-
logical” or “man-environment theme” as one amongst several others.3 However, the
history of the discipline shows that this “ecological” man-earth theme constituted –
at least latently – the organising centre and actual legitimatory foundation for all
important geographical research programmes from the 18th through to the 20th
century.4 It may have taken more of a background role due to the disciplinary breaks
that occurred in the second half of the twentieth century, but it has by no means lost
its influence.
From the very beginning, geographical research and literature consisted in
regional or worldwide (“comparative”) applications of this basic idea. Vast amounts
of material from all manner of scientific and popular literature were dealt with in the
light of the man-earth theme. The “ecological theme” determined not only the foun-
dations and structure of regional geography but also the way in which geography as
a whole constituted its objects. Whether consciously or not, even the Physical
Geographers described the nature of the earth in such a way that their descriptions
could still be linked to the man-earth theme.5 Even today, many geographers and their
scientific statesmen in particular (at least in the German-speaking world and when
they are up on their soap-boxes) are convinced that the essential nature of “geogra-
phy” and its inherent power make it a comprehensive ecology – indeed, make it the
ecology and environmental science per se. But what do they mean by “ecology”?
Where did this geographical ecology come from and what has become of it?
Geography as “Cultural Ecology”
Outside geographical circles, in cultural anthropology and ethnology, for example,
the core paradigm of classical geography described above is characterised as
“cultural ecology” or “the cultural ecological perspective” and, for some consider-
able time now, has been described roughly as follows: A comparative analysis,
carried out between very different environments and cultural forms, of the relation-
ships/interactions between the physico-biotic environment of human populations
on the one hand, and their modes of behaviour, social organisation and culture on
the other. “Culture” here was – and is – taken to mean not only the material culture
3 In the German-language literature, Hard 1973 (pp. 79 ff.) and Bartels and Hard 1975 (p. 90)
placed even greater emphasis on the diachronic and synchronic heterogeneity of geography, and
described in detail each of the many research perspectives contained within this “diffuse
discipline”.
4 Cf. Eisel 1980. A two-part anthology of geographical responses to the question of what geogra-
phy is also provides impressive confirmation of this with regard to the self-perception of geogra-
phers from the 18th century up to the present day (Schultz 2003).
5 For geomorphology see Böttcher 1979.
35325 Geography as Ecology
or the “technical culture” by means of which humans cope with their environment,
but rather every aspect of society and culture that appears to be affected in any way
by this process of “coping with the environment”.6 These and other descriptions of
“cultural ecology” are a very accurate rendering of the aforementioned core para-
digm of geography in the modern era, too; even the frequently quoted phrase intro-
duced by the “founding father” of modern university geography – “the earth as
mankind’s dwelling and place of instruction”7 – essentially had this cultural-ecological
meaning and was consistently understood as such even after Ritter’s death.
From time to time since then, even geography itself has explicitly defined its
essence in terms of “ecology”, as for example in American geography between
about 1910 and 1930 (thereby establishing a clear link to the contemporary
American plant ecology of the time). One example of this is the statement:
“Geography is the science of human ecology”, i.e. “the science of the mutual rela-
tions between man and his natural environment” (Barrows 1923, p. 3). The object
of geography, however, was not to be seen as the “natural environment” as such, but
rather its “habitat value” – in other words, the natural environment as an object of
perception, judgment and utilisation by man. Thus, this “geography as human ecol-
ogy” was also a version of the “Rittersche Wissenschaft”.8
Geography as a Regional and Regionalising Cultural Ecology
Geographical attention was directed above all at the level of landscapes and regions.
The whole logic of the programme described above was particularly well tailored
to pre-industrial (especially peasant) life worlds, as long as they did not seem to
have been significantly transformed (yet) by the global market, industrialisation
and other kinds of modernisation and globalisation. Of course, themes such as
industrialisation and urbanisation increasingly had to be addressed in geography,
too, but it was now more difficult to establish a link between this modern world and
the man-nature theme.
6 For example, Steward 1955, pp. 40 f.; for further references, see Krewer and Eckensberger
(1990). The term “cultural ecology” is used in a similar way in English-language geography as
well (cf. e.g. Johnston 1988), while in German-speaking geography the term “human ecology”
seems to have become established, cf. the recent anthology by Meusburger and Schwan (2003).
7 “die Erde als Wohn- und Erziehungshaus des Menschengeschlechts” (Herder 1784–1791, 1966
edition; pp. 59–67).
8 For more detail about this link to biological ecology from a history of science perspective, cf.
Fuchs 1966, 1967. Around 1920, Chicago – an important location in the history of American
geography – was not only a stronghold of plant ecology (see Chap. 20) it was also the stronghold
of a variety of sociological urban research which thought largely in terms of ecological analogies.
In this famous urban sociology of Chicago, however, “human ecology” (later to become “social
ecology”) usually meant something different from what it meant in the geography of the time,
namely the study of the “social environment” of human populations. On the reception of this
sociological “human” or “social” ecology in German urban research studies from the 1970s
onwards see, amongst others, Hamm 1990.
354 G. Hard
The bulk of interest, moreover, was directed less towards individual behaviour
than towards the relation between entire (groups of) life forms or entire cultures and
their regionally differentiated “natural” or physico-biotic environments. Since the
19th century, therefore, geographical literature in every European language has
provided an inexhaustible repertoire of original case studies and overviews extending
across the entire globe.9 One literary highlight in this field, indeed one highlight of
classical geography as a whole, is the géographie humaine of Vidal de la Blache
(1845–1918) and his school (“l`école française de géographie”), which remained
dominant in French universities and schools until well into the mid-twentieth century,
and which continues to define the general image of geography even today.10
In this geographical cultural ecology an extraordinary spatial perspective has
emerged. Rather than looking at nature, natural conditions and culture as a whole,
it was the spatial “natural plan” and the spatial “cultural plan” in particular (or even