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1
A Biased History of Community Ecology Community
Ecology, Fall 2006 T. E. Miller Dept. of
Biological Science Florida State University
2

• Pre 1900. Cataloging a Balanced Nature
• 1900-1920. The Americans and Plant Succession
• 1920-1950. Animal Population Biology
• A Digression to Ecosystem Ecology
• 1950-1978. Competition and the niche
• 1979- present. Anarchy and new directions
• What can we learn from our history?

3
Pre-1900 -- Cataloging a Balanced Nature
Some claim that the father of ecology is
Gilbert White (1720-1793) who wrote The Natural
History of Selbourne (1789). Whites view of the
world as being in some kind of natural balance
was a precursor to the superorganism ideas of
Clements and others.
'Earthworms, though in appearance a small and
despicable link in the chain of nature, yet, if
lost, would make a lamentable chasm. Worms seem
to be the great promoters of vegetation, which
would proceed but lamely without them.' 'A vast
rain. The hay lies about in a miserable way.'
4
Alexander von Humboldt (1773-1858) was one of the
early catalogers of vegetation. Others include
Raunkiaer, Kerner, de Candolle, and
Warming. Humboldts expeditions to South America
collected 60,000 specimens, 3,000 species new to
science, all presented in detail in 17 lavishly
illustrated volumes.
Von Humboldts camp along the Orinoco
Cinchona condaminea (Quinine) from Plantes
equinoxiales
5
The term father of ecology gets thrown around a
lot (by the way, the position of mother of
ecology seems to still be open). After Gilbert
White, another candidate is Johannes Warming
(1841-1924), who studied woodland savanna in
Brazil for 3 years. Then 30 years later he wrote
theOecologia of Plants, which was a model for
later ecological studies. He also organized one
of the first formal courses in ecology, covering
plant morphology, physiology, taxonomy, and
biogeography.
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Two legacies from this pre-1900 history 1. The
balanced, equilibrium community -- Whites view
of an organized nature lead to a later view of
communities as organized systems with emergent
properties. 2. Ecology as cataloging
species -- often correlated species
abundances with physical factors such a soil
and climate. -- associated with plant
physiology.
These could either be viewed as legacies or
leg-irons.
7
The view that communities were balanced and at
equilibrium was common throughout writings at
this time. Stability was a common view in many
sciences at the time. Herbert Spencer was an
19th century philosopher who was a great defender
of Darwins ideas. His writing was described as
being obscure and overly abstract, but he is
thought to have originally coined the term
survival of the fittest. His very influential
book, First Principles (1862), discussed
similarities between organisms, machines, and
society.
8
When we see that in a mammal, arresting the
lungs quickly brings the heart to a stand that
if the stomach fails absolutely in its office all
other parts by-and-by cease to act that
paralysis of its limbs entails on the body at
large death from want of food, or inability to
escape that loss of even such small organs as
the eyes, deprives the rest of a service
essential to their preservation we cannot but
admit that mutual dependence of parts is an
essential characteristic. And when, in a society,
we see that the workers in iron stop if the
miners do not supply materials that makers of
clothes cannot carry on their business in the
absence of those who spin and weave textile
fabrics that the manufacturing community will
cease to act unless the food-producing and
food-distributing agencies are acting that the
controlling powers, governments, bureaux,
judicial officers, police, must fail to keep
order when the necessaries of life are not
supplied to them by the parts kept in order we
are obliged to say that this mutual dependence of
parts is similarly rigorous. Unlike as the two
kinds of aggregates otherwise are, they are
alike in respect of this fundamental character,
and the characters implied by it. Spencer 1897
I-2452-453
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Ecology as Cataloging Species Remember that
Alexander von Humbolt and Aime Bonpland,
published Essai sur la geographie des plantes in
1807, in which they described vegetation based on
climate, elevation, latitude, and established
'growth form' concept.
This approach reached its zenith with Josias
Braun-Blanquet (1884 - 1980), who developed
methods of community sampling, as well as
analyses methods of data reduction and
association. This approach became known as the
Zurich Montpellier School of Phytosociology or,
sometimes, the Braun-Blanquet system. The
Braun-Blanquet method of plant community
classification is still used today, primarily in
Europe
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Plots are placed in areas representative of the
vegetation of the entire stand. Plot data are
then compiled into tables (species by plots), the
species sorted to identify those that co-occur in
certain patterns, and then stands grouped into
associations. The associations can then be
compared to one another to determine which groups
of species best exemplify the association, either
by being dominant or restricted to the
association.
Josias Braun-Blanquet and wife (1884 - 1980).
In England, Arthur Tansley (1871-1955) utilized
this approach while coordinating a large project
to map the vegetation of the British Isles,
published in 1911. He later worked to show how
the vegetation was affected by soil, climate and
the presence of wild and domesticated animals.
Tansley was also instrumental in starting one of
the first professional societies in ecology, the
British Ecological Society in 1913 (ESA was
started in 1911), and he coined the term
ecosystem.
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Americans also took to cataloging species,
resulting in some of the great collections that
we have today. Among them was Luis Agassiz,
Swiss naturalist and founder of the Museum of
Comparative Anatomy at Harvard. We obviously
still catalog species and investigate species
associations as an important first step in
community ecology. Larger scale pattern analysis
is also at the center of Landscape Ecology and
now GIS approaches.
This statue of Louis Agassiz was toppled from its
niche above the arches at Stanford University by
the San Francisco earthquake, Apr. 18, 1906,
magnitude 8.3.
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1900-1920 The Americans and Plant Succession
Stephen Forbes (1844-1930) is considered the
father of American Ecology. He fought in the
Civil War, but later established a seminal
research program at the University of Illinois.
He is best known for his early studies on food
chain relationships. "These fluviatile lakes
are most important breeding grounds and
reservoirs of life, especially as they are
protected from the filth and poison of towns and
manufactories by which the running waters of the
state are yearly more deeply defiled. The Lake
as a Microcosm, 1887
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"Nowhere can one see more clearly illustrated
what may be called the sensibility of such an
organic complex, expressed by the fact that
whatever affects any species belonging to it,
must have its influence of some sort upon the
whole assemblage. He will thus be made to see
the impossibility of studying completely any form
out of relation to the other forms the
necessity for taking a comprehensive survey of
the whole as a condition to a satisfactory
understanding of any part. If one wishes to
become acquainted with the black bass, for
example, he will learn but little if he limits
himself to that species. He must evidently study
also the species upon which it depends for its
existence, and the various conditions upon which
these depend. He must likewise study the species
with which it comes in competition, and the
entire system of conditions affecting their
prosperity and by the time he has studied all
these sufficiently he will find that he has run
through the whole complicated mechanism of the
aquatic life of the locality, both animal and
vegetable, of which his species forms but a
single element." Stephen Forbes, The Lake as a
Microcosm, 1887
14
Henry Chandler Cowles (1869-1939) was a former
geologist who became a botanist at the University
of Chicago. He, with others, shifted plant
ecology to thinking about succession -- the
regular progression of communities through a
series if stages to some climax community. He
produced a large number of influential students,
including Henry Gleason, William Cooper, Edgar
Transeau, and Emma Lucy Braun and he helped found
the Ecological Society of America (1911).
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Cowles classic work was on sand dunes of Lake
Michigan. He argued that a time-series of plant
communities could be seen as one moved inland
from the open lake beach across ancient
shorelines through the shifting dunes to the
interior forest. Along this route, scrubby beach
grass would give way to flowers and more
substantial woody plants, cottonwoods and pines
would be seen yielding to oaks and hickories, and
one would finally encounter the climax forest of
beeches and maples.
16
A former student of Cowles recounts taking the
train to the Indiana Dunes in long dresses, then
taking off the dresses at the station to reveal
riding britches and boots.
17
Fredrick Clements (1874-1945) was perhaps the
most famous of all the early American ecologists,
although his is generally known for concepts that
are now thoroughly rejected. In particular, he
is known for promoting the view of communities as
superorganisms.
18
Clements traveled all over the western United
States, where he developed methods of vegetation
sampling and described plant growth strategies
for a variety of biomes. His superorganism
view was that plant communities could be viewed
as organisms that followed prescribed
life-cycles to some climax community. Each
species in the community contributed to the
inherent properties of the organism. This
view was immediately disputed by many plant
ecologists, most strongly by Henry Gleason.
19
Henry Gleason (1882-1975) was a student of Cowles
who argued against the superorganism concept and
presented an alternate view. He recognized
that most plant species occur along environmental
gradients and suggested that each species was
responding individually to the factors such as
water, light, or temperature, rather than as any
organized system. This is the individualistic
concept.
20
"Succession is an extraordinarily mobile
phenomenon whose processes are not to be stated
as fixed laws, but only as general principles of
exceedingly broad nature and whose results need
not and frequently do not ensue in any definitely
predictable way" H. A. Gleason, The structure
and development of the plant association,
1917 Gleason was considered to be somewhat of a
heretic until his ideas were picked up again in
the 1940s.
21
The historian Sharon Kingsland has a new view of
the history of ecology which I suspect will not
be widely accepted. She puts the history of
plant ecology in the context of the economic
expansion of the country and the need for science
to support that expansion. She suggests that the
New York Botanical Garden was the "midwife for
the emergence of ecology in the United States,
in part because of its connections with Fredrick
Clements and the Carnegie Institute Desert
Laboratory, outside of Tucson, Arizona.
Clements superorganism concept and deterministic
view of plant succession held sway for almost 30
years, too long in the view of many plant
ecologists. In the 1950s and thereafter, very
different approaches -- based on early work of
Henry Gleason but developed by the Wisconsin
group of plant ecologists and Robert Whittaker --
took hold. These approaches viewed vegetation as
a continuum in relation to changing climatic and
other factors. Clements's views were repudiated
at the 75th-anniversary meeting (1990) of the
Ecological Society of America, a panel of eminent
ecologists could find nothing good to say about
Clements except that he did do experiments.
Paraphrased from a book review by Nancy Slack,
Science 2, February 24, 2006.
22
Naturalists and the Balance of Nature (pre 1800)
Phytosociology and Collectors (1800-1900)
Plant Community Ecologists and Succession
(1900-1920)
23
1920-1950. Animal Population Biology
Not much happened in animal community ecology up
until 1920 or so. Of course, Stephen Forbes had
written about food webs, but Gleason wrote a 25
year history of ecology without even mentioning
animals. The emphasis in animal ecology at the
time was on the individual and, later, the
population, rather than the community. Folks
such as Darwin, Huxley, and Agassiz used the
approach of observing, dissecting, and comparing
individual anatomy and behavior. And, contrary to
the plant ecologists, some evolutionary thought
was included in animal ecology. But, animal
ecologists generally did not concern themselves
with patterns of association or diversity.
24
Charles Elton (1900-1991) was an animal ecologist
who wrote two important books during this time,
Animal Ecology (1927) and The Ecology of
Invasions by Plants and Animals (1941). He
exemplified the thinking of many animal
ecologists at the time that the controlling
processes of communities were species
interactions. He presented interactions webs of
communities, suggesting that the action of
competition and predation was structuring
communities.
Elton is also the source for one of the two major
definitions of the niche. He wrote, "When an
ecologist says 'there goes a badger' he should
include in his thoughts some definite idea of the
animal's place in the community to which it
belongs, just as if he had said 'there goes the
vicar'. Charles Elton, by the way, has been
called the father of invasion ecology.
25
Alfred Lotka, chemist, demographer, ecologist and
mathematician, was born in Austria( actually, now
part of Ukraine). He came to the United States
in 1902 and wrote a number of early theoretical
articles on chemical oscillations, and then
authored a book on theoretical biology (1925). He
may be best known for the predator-prey model he
proposed, at the same time but independent from
Volterra (hence, the Lotka-Volterra model). He
then left (academic) science and spent the
majority of his working life at an insurance
company (Metropolitan Life). In that capacity he
became president of the PAA (the Population
Association of America). He did propose that
evolution maximized energy flow, a concept that
Howard Odum later applied to ecosystem ecology.
26
Georgyi Gause (1920-1986) was a Russian
microbiologist who was professor at the
Timiriasev Institute in Moscow, then later the
director of the New Antiobiotics Research
Institute. He studied the population structure
bacteria and protozoa and, in 1932, published
what has become known as the competitive
exclusion principle. He demonstrated this
principle using experimental work on protozoa in
culture much of his data has been used by
ecologists since.. In 1934, he published The
Struggle for Existence, a groundbreaking work on
the mathematics of population growth.
27
Victor Shelford, a student of Clements, applied
the ideas being used in plant community studies
to investigate tiger beetles in the 1940s.
28
A Digression on Ecosystem Ecology
Many trace ecosystem ecology back to Eltons 1927
Animal Ecology. In this book, he discusses food
webs and energy transfer between trophic levels
and, in particular, proposed pyramids of the
number of individuals or biomass found at
different trophic levels. This stimulated a
group of ecologist to consider how energy may
contribute to this pyramid-like pattern, based on
the energy conserved and lost at each trophic
level.
Lo! Big Fshes are eating Small Fishes. That I
knew for a Long Time - a work by Peter Brughel
29
George Evelyn Hutchinson (1903-1991) was born and
raised in Cambridge, then took a position in
South Africa. He later moved to a postdoctoral
position at Yale where he stayed. He was the
premier limnologist of his time and fostered a
number of influential students, including Howard
Odum, Larry Slobodkin, Robert MacArthur, and Fred
Smith. He could turn a phrase, with titles such
as Homage to Santa Rosalia or Why are there so
many animals and The Ecological Theatre and
Evolutionary Play.
father of modern limnology
30
Raymond Lindeman received his Ph.D. in March,
1941 from the University of Minnesota, and began
postdoctoral work with G. Evelyn Hutchinson at
Yale that September. His career was cut short by
his death in April, 1942 at the age of 27. The
paper for which he is most remembered was
published posthumously in 1942 (The
trophic-dynamic aspect of ecology, Ecology 23
399-418). The paper is the result of his thesis
work on Cedar Creek Bog, Minnesota he already
had a draft version completed when he joined
Hutchinson's lab. Hutchinson was instrumental in
getting the paper accepted for publication (it
was initially rejected by reviewers!). The
editor, Thomas Park, eventually published it,
saying Time is a greater sifter of these matters
and it alone will judge the question. It may be
the single most influential paper in ecology.
31
Eugene Odum
Eugene Odum (University of Georgia) and Howard
(Tom) Odum (University of Florida, where very
influential in the establishment of ecosystem
ecology as a discrete field
32
Hubbard Brook experimental watershed.
The Hubbard Brook Experimental Forest is a 7600
acre tract which allows monitoring and
experiments at the level of entire watersheds.
Long term studies begun by Gene Lichens and
Herbert Bormann (Dartmouth) have been conducted
since 1955. A large experimental study in which
all vegetation was removed from an entire
watershed was initiated in 1963. It is also well
known for some of the earliest American studies
on acid rain, especially a critical 1968 study.
33
Naturalists and the Balance of Nature (pre 1800)
Phytosociology and Collectors (1800-1900)
Plant Community Ecologists and Succession
(1900-1920)
Animal Population Biologists and
Theory (1920-1950)
The Niche and the Rise of Competition
34
1950-1978 Species Packing and Niche
Robert Whittaker (1920-1980) was a plant
ecologist who carried on the earlier ideas of
Gleason about communities along gradients. He
contributed significantly to the development of
ordination and gradient analysis, the measurement
of diversity, and the forces of succession. His
own still often cited work primarily examined
vegetation along elevation gradients. His
gradient work simulated him to think about
species diversity at different spatial scale
alpha, beta, and gamma diversity.
35
Back to Evelyn Hutchinson, who definitely wasnt
just a limnologist or ecosystem ecologist. One
of Hutchinsons many contributions was the niche
defined as an n-dimensional hypervolume (1959.
Am. Nat. 93145-159). With several very
influential papers in the 1950s he got ecologist
thinking in a rigorous framework about the
competitive exclusion principle, species packing,
and species diversity.
36
Robert MacArthur (1930-1972) was a student of
Hutchinson, then later a faculty at Princeton,
who influenced ideas on the niche, population
growth and regulation, island biogeography, and
patch dynamics. He has a very long lasting
effect on ecology through his clear development
of simple theory. To do science is to search
for repeated patterns, not simply to accumulate
facts.
MacArthur stimulated experimental work in
community ecology, as well as providing a number
of innovative theoretical concepts. Some say he
misled us down a garden path.
1958. Ecology 39599-619.
37
Robert May was an Australian physicist who took
his Ph.D. degree in superconductivity. He then
became interested in questions of social
responsibility and started investigating patterns
of population growth, resulting in some of the
earliest work on chaotic population growth. May
balked at a statement that complex ecosystems
were more stable than simple ones, eventually
leading to his matrix modeling of communities. He
moved from Sydney to Princeton and, eventually,
to Oxford where he has also served as as the
chief scientific officer to the British
Government. He was knighted in 1996 and made a
life peer in 2001. He currently serves as a
cross-bench member of the House of Lords and is
often referred to as Sir Bob.
38
Recent Major Schools in Community Ecology
1950 -- Yale University -- Princeton
University 1960 -- University of
Michigan 1970 -- Duke University --
University of Washington -- University of
California, Santa Barbara 1980? 1990? 2000?
39
This is a very typical example of niche overlap
work, by Dinah Davidson. Overlap in ant seed use
was used to estimate per-capita interaction
coefficients, which were in turn used to quantify
indirect effects (Davidson 1984??)
40
Bluegill sunfish
Pumpkinseed sunfish
Green sunfish
41
Bluegill sunfish
Pumpkinseed sunfish
Green sunfish
42

• What is the stated purpose of this paper?
• What general approach or methods are used to
  accomplish that purpose?

We report a set of experiments designed to
examine niche shifts in three species of
freshwater sunfishes. . . .
Four (although 18 were advertised) identical
artificial ponds were used. Each species was
grown alone in a single pond in monoculture,
stocked with 900 individuals. The fourth pond
was stocked with 900 individuals of each species,
all thrown in together.
43

• Are the data sufficient and clearly presented?
• What are the primary stated conclusions?
• Is the main question important (not to be
  confused with interesting)?

44

• 6. Go back and compare the stated purpose of the
  paper (1) with the main conclusions (4). Do
  they match?
• Do the data support the conclusions? Would
  another approach have been better? Are the data
  over-interpreted or is the interpretation just
  wrong.

45

• What overall contribution does this paper make to
  community ecology?
• 9. What new questions are raised and what
  further work is suggested by this study, either
  directly or indirectly?

46
1978-present Anarchy and new directions
Starting in 1979, Don Strong, Dan Simberloff,
Larry Abele and graduate students at Florida
State vigorously attacked several paradigms of
the time. In particular, they debunked many
previous studies on character placement and the
importance of competition in structuring
communities and promoted the use of null models
in ecology. For this, they became known as the
Tallahassee mafia. Several other individuals
have a very strong effect at this time, including
Caswell (non-equilibrium coexistence) and
Hurlbert and Underwood (design and statistical
rigor).
47
1. Competition and character displacement
Strong, Sycek, and Simberloff (1979)
re-evaluated data that had previously been used
to support the concept that competition
structured patterns in communities. In the
example below from Peter Grants work on Darwins
finches, bars give actual ratios in beak size
between pairs of species, while circles give the
null expectation. Character displacement would
be indicated by bars above circles. Ultimately,
this led to the development and use of null
models.
48
2. Non-equilibrium approaches The nature in
balance perspective, as well as mathematical
considerations, had led to the assumption that
most populations and communities were at or near
some equilibrium. In population biology, the
equilibrium view gave way to a general
recognition that density-dependent and stochastic
forces interacted to determine species abundances
(Strongs density-vagueness).
In community ecology, a number of different
factors direct us away for equilibrium views.
These include -- patch dynamics (Levins and
Culver, Paine and Levin) and disturbance theory
(Connell) -- non-equilibrium coexistence
(Caswell) of predator and prey -- coexistence of
equal competitors (Hubbell) -- assembly and
chance, lottery hypothesis (Chesson)
49
3. Experimental Rigor Several papers in the
late 1970s and early 1980s noted wide-spread
problems with experimental design in ecological
studies. This included -- the lack of
appropriate null models (Strong, et al.
previously discussed) -- pseudoreplication
problems (Hurlbert 1984 showed that 48 of
studies were psuedoreplicated) -- lack of
appropriate controls and other statistical
problems (Underwood 1981 found 78 of marine
studies that used analysis of variance contained
statistical errors) Ecologists were forced to
shift from being simple interpreters of patterns
of natural history to conducting more rigorous
and formal experiments.
50

• Recent History
• Competition and character displacement
• Non-equilibrium approaches
• Experimental Rigor

The response to 1 was very predictable. First,
came denial, followed by a new emphasis on the
importance of predation. This in turn was
followed by a too much work that argued whether
competition or predation was more important.
Finally, we turned to more integrated work that
investigated the contributions of both forces, as
well as their interactions. This included some
interesting theoretical work by Holt (apparent
competition), as well as Bender et al., Stone,
Yodzis, and Wootten.
51
Top-down vs. bottom-up studies.
52
It has to be pointed out that some scientists
were already doing excellent work in this regard.
This includes Henry Wilbur and his lab working
on amphibians in small ponds and streams, Earl
Werner and his lab working on fish communities in
ponds, Bob Paine and his students working on
predation and disturbance in the intertidal, Joe
Connell working in intertidal systems, as well as
tropical forests, and Jane Lubchenco, also
working in intertidal systems. These folks were
all working in systems where predation was
clearly important, in part for determining the
intensity of competition.
53

• Recent History
• Competition and character displacement
• Non-equilibrium approaches
• Experimental Rigor

The equilibrium assumption (2) had already
bothered many practical ecologists. Although it
could be argued that several earlier papers were
also important, I think three critical papers
were Joe Connells clear explanation of the
intermediate disturbance hypothesis (1978), Hal
Caswells (1978) work on predator-mediated
coexistence in a non-equilibrium system and Steve
Hubbells (1979) non-equilibrium model of
tropical forest diversity (more recently
developed as neutral theory). Later important
work included Chessons lottery model (and
Chesson and Huntleys work) and, eventually
metapopulation and metacommunity approaches.
54

• Recent History
• Competition and character displacement
• Non-equilibrium approaches
• Experimental Rigor

Methodology changed quite a lot after 1978, with
Florida State faculty and students playing a
large role. Null models (see Caswell 1976), in
particular, were promoted by Strong, Simberloff,
Gotelli and Graves of FSU to address questions
about character displacement. The importance of
experiments (hypothesis testing) as opposed to
observational studies (hypothesis creating)
became well accepted. Experimental design became
somewhat more rigorous, addressing concerns about
psuedoreplication and power. Statistical power
on personal computers increased greatly, with had
both positive and negative affects. However,
reviews suggest that many problems still exist.
Psuedoreplication, poor designs, and lack of
power are still problems that frequently appear
in manuscripts as well as published papers.
55
Naturalist and the Balance of Nature
Phytosociology
Plant Community Ecologists and Succession
Animal Population Biologists and Theory
The Niche and the Rise of Competition
Anarchy and Experimental and Statistical Rigor
Spatial and Temporal Heterogeneity -
Metacommunities
56
What Now? I have no idea! One cannot write
history as it happens -- it must write itself as
some passage of time. Some particularly
interesting areas are -- functional
groups -- neutral theory (well, maybe not . .
.) -- metacommunities (but, not particularly
new) -- evolution in a community context ???
57
P. WERNER
MILLER
WINN
Sprugel, D. G. 1980. A "pedagogical genealogy" of
American plant ecologists. Bull. Ecol. Soc. Amer.
61197-200.