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Graham Harris
Chapter
39
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40 The complexity of ecology
Intellectual foundations
Many others have commented that it was Darwin’s fundamental insights about
the struggle for existence that began the scientific study of ecology. Indeed,
chapter 3 of Darwin’s Origin of Species on the struggle for existence, and his
concluding remarks about a ‘tangled bank’, are what would now be regarded
as straightforward ecology:4 population interactions, community structure and
competition.
In Darwin’s era there was a firm belief in the concept of plenitude and of the
Great Chain of Being. These ideas have deep roots in the concept of the spiritual
unity of the universe. Plenitude defines the desirable (and perfect) state as the
one with the greatest richness of species; in short, the world is brim-full of life
and the struggle for existence is continuous. In addition humankind was seen
as the pinnacle of evolution with the Great Chain stretching upwards from the
lowliest bacteria and protozoa all the way to us. The hand of God was seen in
this design. These are very old concepts. Glacken5 traced the roots of western
ecological thinking back long before Darwin to Greek philosophy and showed
how, both then and now, ecological thinking is strongly influenced by arguments
from design. Glacken wrote
Following Glacken’s logic, Marshall7 pointed out that the argument from
design has a number of accompanying assumptions about concepts of uni-
formitarianism, balance, harmony, equilibrium and the concept of life as a
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Intellectual foundations 41
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42 The complexity of ecology
is such a key part of the modern world view that Appleyard’s critique of science
is centred on the ‘severance of knowledge and value’ that he argues has taken
place.12
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Natural resource management 43
markets are so low that farming enterprises around the world are unprofitable,
how do we expect farmers to be environmental managers as well as produc-
tive farmers? How do we therefore manage the ‘rest of nature’? Many countries,
including the USA and the states of the European Union (EU), resort to var-
ious kinds of agricultural subsidies or stewardship payments. Increasingly, as
I shall discuss later, market activity is expanding to encompass commodities
other than agricultural produce to include ecosystem services and conservation
outcomes.15
Sustainable land management requires profitability and resources in order
to make the necessary investments for restoration and conservation. There is
an equal need to manage and restore agricultural and urban landscapes, which
are much more heavily modified than the largely natural landscapes of parks
and reserves. Furthermore, in a non-equilibrium world of emergence and con-
stant change, it is impossible to place a fence around any area and conserve or
protect it for all time: there will always be both externally and internally gen-
erated changes in ecosystems and landscapes. Everywhere is on a trajectory to
somewhere, driven by complex interactions between components. It is therefore
necessary to think clearly about the nature of ‘the rest of nature’, to attempt to
find the underlying rules and mechanisms and to try to conceive practical ways
to manage and restore natural landscapes and ecosystems. This is the challenge
of the twenty-first century: restoring biodiversity and managing biophysical real-
ities through the involvement of the other parts of the ‘systems of systems’. We
cannot ignore the impacts of global markets or the importance of local commu-
nity involvement. This is a complex issue of conflicting beliefs and values, and
biophysical realities.
Restoration of landscapes and rivers is going to be done by local communities.
For example, the Australian dryland salinity problem is going to be fixed by farm-
ers and graziers changing their behaviours and agricultural practices. Yes, there
is a role for government to provide regulation, incentives and some investment;
but the majority of the knowledge, effort and success is going to come from
local groups. The decisions that rural communities take in their management
of environmental and agricultural assets are based on culture and values, and
on perceptions of risk and return. Social capital is critical. Success depends on
having suitably sustainable management options, on learning, innovation, rela-
tionships, networks and cultural change. The difference between environmental
groups and farmers lies in their perceptions of evidence, knowledge, culture,
beliefs and values. There is therefore an urgent need to engage rural and urban
communities -- and their beliefs and value sets -- and to enlist their help in
restoration. This requires the exact opposite of confrontation and oppositional
politics. Oppositional politics is often required to identify and raise the visibility
of an issue and to ‘grab people’s attention’. Once that has been done, kitchen-
table mentoring works better than public criticism.
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44 The complexity of ecology
The only way forward is to recognise the importance of maximising and man-
aging multiple capitals in the landscape and the necessity of not separating the
financial from the natural, the utilitarian from the objectivist, either ethically
or physically. In other words, the entire landscape -- people, biodiversity and
productive enterprises -- must become the unit of management. This concept
involves rejection of ‘special’ wilderness places as separate from the ‘ordinary’
and significantly widens the definition of what constitutes ecology and what con-
stitutes science: this is very much post normal ecology. This, of necessity, involves
a much more complex set of values, cultures and ethics than is presently being
demonstrated by any party; hence the need for ‘lifting the level of the debate’
around possible futures, and the search for alternative pathways to sustainabil-
ity. Because the objectivist ‘greens’ tend to be an urban group in opposition
to the more utilitarian rural populations this also involves capacity building on
both rural and urban communities and the bringing together of a more inclusive
debate. In the end these problems are only going to be solved by offering rural
communities sustainable options (i.e. environmentally sound, socially acceptable
and profitable options) based on justice, equity and fairness. A systems view and
ethical decision making are of paramount importance. The problem with many
environmental and landscape management problems is that we lack sustainable
options -- and this is going to require research and community consultation,
working through a dialogue and an inclusive process. Science and ecology are
necessary but not sufficient. So, also for a number of reasons, conservation is
necessary but not sufficient. The world view, values and sense of place that we
hold have a critical role to play in the way we view and manage the world.
This world view has changed over time and is changing rapidly even as I write.
Ideas and concepts are changing to a more complex and inclusive view, a more
dynamic and challenging view -- a view that I argue is going to be more sustain-
able in the long run, but one that is going to require a considerable change in
thinking.
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World views and ecological concepts 45
Design concerned with overarching conceptual schemes and with balance, har-
mony, equilibrium and ultimate causes. ‘The whole exists first and its design
explains the actions of the parts’,16 whereas the mechanistic approach assumes
that the ‘actions of individual parts of a whole are explained by known laws,
and the whole is the sum of the parts and their interaction’. Basically, these are
‘top-down’ and ‘bottom-up’ approaches, which some argue are incompatible; so is
there a ‘Grand Canyon’ down the middle of our thinking? [Readers can probably
guess by now where this argument is heading -- the bounded SGC view builds
the bridge . . . but let us review history for a while before we come to that.]
Hengeveld and Walter17 would argue for the Grand Canyon effect. The devel-
opment of ecological thought and theory must be seen in the context of the
longer-term (sometimes cyclical) developments in society at large. From its roots
in the eighteenth and nineteenth centuries, ecology has always been part and
parcel of the broader development of philosophy and epistemology. The theor-
etical development of the ‘organic’ approach comes from Darwin and from argu-
ments from design. Darwin adapted an idea from Adam Smith that equilibrium
could be achieved by free competition at the community level, and this set the
foundation of the very common ecological principle that almost everything is
explicable in terms of competition for common resources. ‘Overall, he effectively
transposed a complex of ideas associated with the human progressionist society
to the natural environment of species, and thus settled a role for competition
at the core of ecology and evolution.’18 In the 1920s and 1930s Lotka,19 Gause,20
Nicholson and Bailey,21 and Fisher22 turned competition into the structuring
force in ecological and evolutionary theory through mathematical theory. By
the 1940s this was developing in conjunction with the ‘modern synthesis’: a
conceptual and mathematical fusion of population genetics, evolutionary the-
ory and ecology.23 (Note that there is no mention of bounded or constrained
solutions to ecological problems.)
Equilibrium population densities in animals were assumed to be reached
through density-compensating mechanisms, boosting birth rates and minimis-
ing death rates at low densities and doing the reverse at high densities. Competi-
tion was seen as a key ‘governor’ of ecological processes, just like the governor on
a steam engine.24 An entire school of population ecology was built around this
assumption.25 Some of the key early proponents were Nicholson and Lack, who
each produced seminal works26 which became popular in the 1950s and much
later. Competition was very much seen as the link between proximate (ecologi-
cal) and ultimate (evolutionary) causes and was enshrined in papers by Hardin
and Hutchinson on the competitive exclusion principle27 and on ‘why are there
so many kinds of animals’28 around 1960. I do not think it is an exaggeration
to say that the idea of competitive equilibrium has been the basis of much, if
not most, ecological theory to this day, particularly in North America. (There
are recent exceptions, which we will come to later.) The equilibrium theory of
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46 The complexity of ecology
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World views and ecological concepts 47
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48 The complexity of ecology
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Bridging the gap 49
us evolution and development in the long run. There is now increasing evidence
that small-scale interactions between populations within patches do structure
the responses to perturbations that we see. Functional traits interact within the
context of environmental factors and gradients.50 So while the overall statistical
patterns of species abundances seem to be more dependent on regional immi-
gration and environmental factors, there are underlying and important local
interactions.
Hengeveld and Walter discuss the strengths and weaknesses of the two
approaches to ecology and point out that, whereas the ‘competitive’ (organic)
approach to ecology has by far the greatest theoretical development, the ‘indi-
vidualistic’ (mechanistic) approach frequently produces data that contradict, or
create difficulties for, the standard theory. The development of theoretical ecol-
ogy has searched for ultimate causes in an abstract form of mathematical theory,
and it has often been divorced from the practical day to day reality and complex-
ity of the natural world. There have been problems applying abstract theory to
the messy world of natural resource and environmental management. Averaging
has often been used to control the ‘noise’ and to extract the patterns of interest.
The data are always a lot ‘noisier’ than the models predict. When I am standing
up to my hocks in someone’s sewage effluent, what does theoretical ecology tell
me? The problem with a theory with progressionist roots is that it does not
recognise the influence of global and regional evolutionary constraints and the
potential for the evolution of ‘neutral zones’ through SGC.
The sheer complexity of the interactions between all the species in the
‘tangled bank’ at a range of scales, the observed variability in space and time
and the limitations of human observers has plagued us from the beginning. The
theoretical (averaged) harmony and order is overlain by much ‘noise’ and vari-
ability, which is reflected in difficulties in field work and data gathering. The
prevalence of dynamic behaviour has meant that ecology has been very good
at developing abstract concepts but rather poor at developing testable theories.
Rob Peters, in his critique of ecology, gave many examples of the problems with
ecological theories.51 It is indeed rather hard to do the usual kinds of scientific
experiment in ecology. Experimental controls, for example, where all else is held
constant while the effects of experimental treatments are examined, are usually
rather hard to come by. Not only is there much small-scale variability in space
and time, but how do we ever find a control for something like an ecosystem
the size, say, of an ocean, a river or a large lake? This is nothing more than the
recognition of the fundamental problems of dealing with uncertainty, indeter-
minism and ignorance, and the unique geological and climatological features
of large parts of the globe. I note here again for future reference that the whole
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50 The complexity of ecology
idea of trying to control ‘noise’ and detect change in ecological data raises some
interesting issues about scientific methods and statistics.
So can we explain where all the ‘noise’ comes from, and is there a way to
go about bridging the ‘Grand Canyon’? Standing in the now dammed and regu-
lated Colorado River, is it possible to find a way out of the chasm? One way, it
seems to me, is to recognise the emergent complexity of the pandemonium of
species and interactions and to recognise that there are issues of connectivity
and context and the possibility of global and regional constraints. In short, the
middle ground is the world of dynamic interactions within and between local
patches, with emergent context at a range of scales.
notes
1. P. Taylor. Unruly Complexity: Ecology, Interpretation, Engagement. (Chicago, IL: Chicago University
Press, 2005).
2. R. P. McIntosh. The Background of Ecology, Concept and Theory. (Cambridge: Cambridge University
Press, 1985).
3. Quoted in McIntosh, The Background of Ecology, pp. 7--8 (see Note 2) and sourced from the
translation in the frontispiece of W. C. Allee et al. Principles of Animal Ecology. (Philadelphia,
PA: Saunders, 1949).
4. See C. Darwin, The Origin of Species, chapter 3, p. 57 and the beginning of the last paragraph
of the book, p. 504 in the reprint of the sixth edition. (New York: The Home Library, Burt
and Co., undated).
5. C. J. Glacken. Traces on the Rhodian Shore. (San Francisco, CA: University of California Press,
1967).
6. Glacken. Traces on the Rhodian Shore, p. 423. (See Note 5.) These ideas are, of course, strongly
present in the widely popular ideas of Intelligent Design.
7. A. Marshall. A postmodern natural history of the world: eviscerating the GUTs from ecology
and environmentalism. Studies in the History and Philosophy of Science: Part C, 29 (1998), 137--64.
8. Glacken, Traces on the Rhodian Shore, p. 708. (See Note 5.)
9. J. Lovelock. Gaia, a New Look at Life on Earth. (Oxford: Oxford University Press. 1979). James
Lovelock. The Ages of Gaia. (Oxford: Oxford University Press, 1988).
10. A. Marshall. The Unity of Nature. (London: London Imperial College Press, 2002).
11. C. Spretnak. The Spiritual Dimension of Green Politics. (Santa Fe, CA: Bear, 1986).
12. B. Appleyard. Understanding the Present. (London: London Picador, 1992); p. 113 in a chapter
entitled ‘From scientific horror to the green solution’.
13. J. R. Saul. Voltaire’s Bastards. (Toronto: Viking, Penguin Books, 1992).
14. S. A. Hajkowicz and M. D. Young (eds). Value of Returns to Land and Water and Costs of Degra-
dation. Final report to the National Land and Water Resources Audit. Project 6.1. (Canberra:
CSIRO Land and Water, 2002). Available from www.nlwra.gov.au.
15. H. A. Mooney, A. Cropper and W. Reid. Confronting the human dilemma. Nature, 434 (2005),
561--562.
16. McIntosh, The Background of Ecology. p. 13. (See Note 2.)
17. R. Hengeveld and G. H. Walter. The two coexisting ecological paradigms. Acta Biotheoretica, 47
(1999), 141--70. G. H. Walter and R. Hengeveld. The structure of the two ecological paradigms.
Acta Biotheoretica, 48 (2000), 15--46.
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Notes 51
18. Hengeveld and Walter. The two coexisting ecological paradigms, p. 144. (See Note 17.)
19. A. J. Lotka. Elements of Physical Biology. (Baltimore, MD: Williams and Wilkins, 1925).
20. G. F. Gause. The Struggle for Existence. (Baltimore, MD: Williams and Wilkins, 1934).
21. A. J. Nicholson and V. A. Bailey. The balance of animal populations. Proceedings of the Zoological
Society of London, Part 3 (1935), 551--98.
22. R. A. Fisher. The Genetical Theory of Natural Selection. (London: Constable, 1930).
23. J. S. Huxley. Evolution: the Modern Synthesis. (London: George Allen and Unwin, 1942).
24. A. J. Nicholson. The role of competition in determining animal populations. Journal of the
Council of Scientific and Industrial Research, 10 (1937), 101--6.
25. S. Kingsland. Modelling Nature: Episodes in the History of Population Ecology. (Chicago, IL: Chicago
University Press, 1985).
26. A. J. Nicholson. The balance of animal populations. Journal of Animal Ecology, 2 (1933),
132--78. David Lack. The Natural Regulation of Animal Numbers. (Oxford: Oxford University
Press, 1954).
27. G. Hardin. The competitive exclusion principle. Science, 131 (1960), 1292--7.
28. G. E. Hutchinson. Homage to Santa Rosalia: or why are there so many kinds of animals?
American Naturalist, 93 (1959), 145--59.
29. D. Tilman. Resource Competition and Community Structure. Princeton Monographs in Population
Biology, 17. (Princeton, NJ: Princeton University Press, 1982).
30. R. H. MacArthur and E. O. Wilson. The Theory of Island Biogeography. Princeton Monographs
in Population Biology, 1. (Princeton, NJ: Princeton University Press, 1967).
31. R. M. May. Stability and Complexity in Model Ecosystems. Princeton Monographs in Population
Biology, 6. (Princeton, NJ: Princeton University Press, 1973).
32. A. Wagner. Robustness and Evolvability in Living Systems. (Princeton, NJ: Princeton University
Press, 2005).
33. H. C. Cowles. The causes of vegetative cycles. Botanical Gazette, 51 (1911), 161--83. F. E.
Clements. Plant succession: an analysis of the development of vegetation. Publication No
242. (Washington: Carnegie Institute of Washington, 1916).
34. E. P. Odum. Fundamentals of Ecology, 3rd edn. (Philadelphia, PA: Saunders, 1971).
35. A. Marshall. A postmodern natural history of the world: eviscerating the GUTs from ecol-
ogy and environmentalism. Studies in the History and Philosophy of Science: Part C, 29 (1998),
137--64.
36. H. A. Gleason. The individualistic concept of the plant association. Bulletin of the Torrey Botan-
ical Club, 53 (1926), 1--20.
37. G. E. Hutchinson. The paradox of the plankton. American Naturalist, 95 (1961), 137--45.
38. G. P. Harris. Phytoplankton Ecology. (London: Chapman and Hall, 1986). Marshall pointed out
that I had taken a Gleasonian view of ecology in this work.
39. U. Sommer (ed.) Plankton Ecology, Succession in Plankton Communities. (Berlin: Springer, 1989).
40. See, for example, C. S. Reynolds. The Ecology of Phytoplankton. (Cambridge: Cambridge Univer-
sity Press, 1984).
41. H. G. Andrewartha and L. C. Birch. The Distribution and Abundance of Animals. (Chicago, IL:
Chicago University Press, 1954).
42. J. Wu and O. L. Loucks. From balance of nature to hierarchical patch dynamics: a paradigm
shift in ecology. Quarterly Review of Biology, 70 (1995), 439--66.
43. J. Ludwig et al. Landscape Ecology, Function and Management. (Collingwood, Victoria: CSIRO
Publishing, 1997).
44. K. J. Gaston. Global patterns in biodiversity. Nature, 405 (2000), 220--7.
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52 The complexity of ecology
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