Robert:Leo' Smith-

West Virginia University

Thomas M. Smith
University ofVirginia
Illustrated by
Robt?rt Leo Smith, Jr.
..
An imprint of Addison Wesley Longman, Inc.
San Francisco Boston New York
Capetown Hong Kong London Madrid Mexico City._ ..
Montreal Munich Paris Singapore Sydney---Tok:yG--Toronto ..
. ~ ;
2
c
Ecology: Meaning and Scope
OUTLINE
Ecology Defined
The Development of Ecology
Plant Ecology
Animal Ecology
Physiological Ecology
Population Ecology
Ecosystem Ecology
Long-Term Ecological Research
Tensions Within Ecology
Plant Versus Animal Ecology
Organismal Versus
Individualistic Ecology
Holism Versus Reductionism
Applied Ecology
Ecology: An Empirical and Experi-
mental Science
Experimental Approach
Field Experiments
Hypothesis Testing
Models and Predictions
. Validation
Summary
Review Questions
CONCEPTS
1. Eqology is the study of the structure and function
of nature.
2. Ecology developed from many ro"ots, but its beginnings
, trace back to natural history and plant geography.
3. Ecology has branched into many subdivisions, many
of them specialized.
4. The principles of ecology provide the scientific basis for
the solution to many of our environmental problems.
5. Modern ecology is an empirical, experimental science.
6 .. Science involves the testing of hypotheses.
7. A hypothesis is a statement that can be tested.
8. Testing hypotheses involves the collection of data by
observation and experimentation, and the analysis of
that data.
9 .. Useful in ecological studies are models, explicit sets
of hypotheses relating pattern and process.
ECOLOGY DEFINED
What is ecology? Ask nonecologists and they will probably
answer that ecology has something to do with the environment
or with saving it. Before the 1960s, few of them could have
given you any answer. If you had asked a biologist in the same
time period, you would probably have gotten some vague
answer implying that ecology was "quantified natural history."
Ecology only became a household word in the 1960s, through
the environmental movement, and then its popular meaning
became confounded with environmentalism.
The origin of the word ecology is the Greek oikos, mean;,
ing "household;' "home;' place to live:' It is derived from
the same root word as economics, "management of the house-
hold." Ecology, then, could be considered as the economics of
nature. Clearly, ecology deals with the organism and its place
to live, its environment. Ecologists continue to work toward a
deeper definition. Here is a sampling of their attempts:
"the study of structure and function of nature." (Odum
1971:3)
"the scientific study of the distribution and abun-
dance of animals." (Anarewartha 1961:10)
"the scientific study of the interactions that deter-
mine the distribution and abundance of organisms."
(Krebs 1985:4)
"the scientific study of the relationships between
organisms and their environments." (McNaughton and
Wolfe 1979:1)
"the study of the relationships between organisms
and the totality of the physical and biological factors
affecting them or influenced by them." (Pianka 1988:4)
"the study of the adaptation of organisms to their
environment." (Emlen 1973:1)
"the study of the principles which govern temporal
and spatial patterns for assemblages of organisms."
(Fencbel1987:12) 1
"the study of the patterns of natUre an4 how those
patterns came to be, and how they change i? space and
time." (Kingsland 1985:1)
"the study of organisms and their environment-and
the interrelationships between the two." (Putman and
Wratten 1984:13)
"the study of the relationship between orgauisms and
their physical and biological environments." (Ehrlich and i
Rougbgarden 1987:3)
The author of the teim, Ernst Haeckel, defined ecology
as "the body of knowledge concerning the economy of
nature-the investigation of the total relationships of the ani-
mal both to its inorganic and its organic environment; includ-
ing, above all, its friendly and inimical relations with those
animals and plants with which it comes directly or indirectly
into contact-in a word ecology is the study of all those com-
plex interrelations referred to by Darwin as the conditions for
the struggle for existence."
None of these definitions is fully satisfactory. They are
restrictive or too vague, and the original definition
----relates only to animal ecology. Most definitions apply to pop-
Chapter 1 Ecology: Meaning and Scope 3
ulation ecology and overlook ecosystem function.
ject has outgrown them.
Instead of trying to define ecology in a manner that
attempts to encompass all that it involves, we might better to
consider a of ecology. works
at characterizing the patterns seen in nature, studying the
complex interactions among organisms and their environ-
ments, and understanditig the mechanisms involved in bio-
logical diversity.
THE DEVELOPMENT
OF ECOLOGY
/
Just as there is no consensus on the definition of ecology, so
there is no agreement on its beginnings. It is more like
a multistemmed bush than a tree with a single trunk.
historians trace the beginnings of ecology to Darwin, Thoreau,
and Haeckel; others to the Greek scholar Theophrastus, a
friend and associate of Aristotle, who wrote about the inter-
relationships between organisms and the environment.
Plant Ecology
The modem impetus to ecology came from the plant geogra-
phers. They discovered that, although plants differed in var-
ious parts of the world, certain similarities and differences
demanded explanation. They looked to climate as a possible
answer, because similar climates supported similar vegeta-
tion. Carl Ludwig Wtlldenow (1765-1812), one of the early
.. iluential plant geographers, championed this explanation.
His ideas caught the attention of a wealthy young Prussian
naturalist, Friedrich Heinrich Alexander von Humboldt
(1769-1859) (Figure l.la). He sailed in 1799 with the French
botanist Aime Bonpland for a five-year expedition through
tro0pical Spanish America. They traveled though Mexico,
.
Cuba, Venezuela, and Peru and explored the Orinoco and
Amazon rivers. Humboldt described these travels and trop-
ical America's plant and animal life in a 30-volume work,
Voyage to the Equatori_al Regions. In these books Humboldt
described vegetation in terms of physiognomy, correlated
vegetation types with environmental characteristics, and
coined the term association.
Travel to the tropics continued to stimulate plant ecol-
ogists. A Danish botanist, Johannes Warming (1841-1924)
(Figure 1.1 b), spent three years studying tropical vegetation
in Brazil, which he described in a book 30 years later. His
major contribution was the book Plantesamfund: Grundtrak
af den okologiske Plantegeografu, published in 1895, in
which he emphasized the importance of moisture, temper-
ature, and soil in patterns of vegetation. The book greatly
influenced the development of ecology. Following Warm-
ing, another tropical traveler, the German botanist Andreas
Schimper (1856-1901), published Pjlanzengeographie auf
physiologischer Grundlage (1898). In it he explained __ _
4 Part I
(a) (c)
FIGURE 1.1 Plant geographers. (a) Friedrich Heinrich Alexander von Humboldt. (b) Johannes
Warming. (c) F. E. Clements.
regional differences in vegetation as a function of moisture
and temperature. -
Other plant geographers stayed closer to home. Never-
theless they made major advances in plant ecology, especially
the emerging concept of vegetation change over time. Anton
Kerner (1831-1898) was one of these geographers. Com-
missioned to survey the vegetation of eastern Hungary and
Transylvania, he described plant developme1.1t in Plant Life
of the Danube Basin (1863). He pioneered the use of exper-
imental transplant gardens at various elevations in the
TYrolean Alps to study the growth and behavior bf plants
taken from alpine and lowland sites. Later, the Polish botanist
Jozef Paczoski (1864-1941) described how plants modify
their environment by creating microenvironments. Because
his book was published in Slavic, it was belatedly discovered
by ecologists outside the Slavic world. He is now recognized
as the father of Plant sociology or phytosociology.
The study of plant communities developed along sepa-
rate paths in Europe and America. In Europe plant ecologists
concentrated on describing the plant community. Christen
Raunkiaer (1860-1938) of Denmark contributed a scheme of
life form classification and quantitative methods of sampling
the data from which could be treated statistically.
Later Josias Braun-Blanquet developed methods of commu-
nity sampling, \data reduction, and the classification and
nomenclature of plant communities. A. G. Tansley, however,
urged a more experimental approach plant ecology. His
views on ecology and research anticipated by years the type
of ecological studies that emerged in the 1970s.
In America plant ecologists were far more interested in
how plant communities develop. Human settlements were
destroying_ the original forests and grasslands, creating pro-
found changes in vegetation. Interest in these disturbances led
to pioneer studies of the dynamics of vegetation and vegeta-
tion changes over time, or plant succession. A doctoral study
on the succession of plant life on Indiana sand dunes by
H. E. Cowles (1897) established the study of plant succession
as one of the central concepts of modem ecology. A major
leader in these studies and the development of American plant
ecology was F. E. Clements (1874-1945) (Figure Llc). Dog-
matic and convincing, Clements quickly became the major
theorist of plant ecology in America. He promoted the idea of
the community as a superorganism and gave ecology a hier-
archical framework still reflected in modem views of the field.
Animal Ecology
Animal ecology developed later than plant ecology and along
lines divorced from it The beginnings of animal ecology can be
traced to two Europeans, R. Hesse of Germany and Charles
Elton of England. Elton's Animal Ecology (1927) and Hesse's
Tiergeographie auflogischer grundlage (1924), translated into
English as Ecological Animal Geography, strongly influenced
the development of animal ecology in the United States. Charles
Adams and Victor Shelford were two pioneering U.S. animal
ecologists. Adams published the first textbook on animal ecol-
ogy, A Guide to the Study of Animal Ecol9gy (1913). Shelford
wrote Animal Communities in Temperate America (1913).
Shelford gave a new direction to ecology by stressing
the interrelationship of plants and animals. Ecology became
a science of communities. Some earlier European ecologists,
particularly the marine biologist Karl Mobius, had devel-
oped the general concept of the community. In his essay "An
Oyster Bank Is a Biocenose" (1877), Mobius explained that
'
the oyster bank, although dominated by one animal, was
really a complex community of many interdependent organ-
isms. He proposed the word biocenose for such a commu-
nity. The word comes from the Greek, meaning "life having
something in common."
The appearance in 1949 of the encyclopedic Principles
of Animal Ecology by five second-generation ecologists from
the University of Chicago-W. C. Allee, A. E. Emerson,
Thomas Park, Orlando Park, and K. P. Schmidt-pointed the
direction modem ecology was to take. It emphasized feeding
relationships and energy budgets, population dynamics, and
natural selection and evolution. '
Still another area of biology, animal behavior, grafted itS
branch onto ecology. Although Darwin, Wallace, and others
described activities of animals, the formal study of animal
behavior began with George John Romanes (1848-1894), who
introduced the comparative method of studying nonhuman ani-
mals to gain insights into human behavior. His approach
depended largely on inferences; in contrast C. Lloyd Morgan
(1852-1936), ,an English behaviorist, emphasized the use of
direct observation and experiment.
After the early 1900s,)!llimal behavior study developed
along four major lines. One was the study of behavioral mech-
anisms, perceptual and physiological. It became known
behaviorism. A second, more relevant to ecology, was the
study of the function and evolution of behavior, including
comparative physiology, pioneered by J. B. Watson in his
book Behavior: An Introduction to Comparative Psychology
(1914). Comparative psychology evolved into ethology. The
three major founders of ethology were Konrad Lorenz, noted
for his studies of genetically programmed behavior; Niko Tm-
bergen, who developed the scheme of four areas of inquiry
(causation, development, evolution, and function); and Karl
von Frisch, who pioneered studies of bee communication and
behavior. After World War II a third field.of animal behavior,
wedded to ecology, appeared. It was eology,
which investigates the way animals interact their living
and nonliving environments, with a special emphasis on how
that behavior is influenced by natural selection. Behavioral
ecology begot a controversial offspring, sociobiology, pio-
neered by E. 0. Wilson in Sociobiology: The New Synthesis
(1975). Sociobiology, concentrating on field observations of
social groups of animals, applies the principles of evolution-
ary biology to the study of social behavior in animals. It
became controversial when some writers attempted to apply
it to humans.
Physiological Ecology
Physiological ecology, or ecophysiology, is concerned with
the responses of individual organisms to temperature, mois-
ture, light, nutrients, and other factors of the environment.
Early plant physiologists studied photosynthesis and plant
growth, including the influence of environment on growth.
Justus Leibig (1840) investigated the role oflimited supplies
of nutrients on the growth and development of plants and
came up witlrthe-''law of_t!le minimum." F. F. Blackman
Chapter 1 Ecology: Meaning and Scope 5
(1905) extended this idea of limiting factors to include-a-max-
imum-a plant could get too much of a good._, thing. Soon
both plant physiologists and plai:J.t ecologists began to work
out the physiological relationships among plants, climate,
atmosphere, and soil:\As plant physiologists deciphered the
mechanisms of and water relations in plants,
ecophysiologists related these functions to plant distributions
and adaptations. After World War II, the field grew rapidly. .
New instrumentation, modem experimental techniques, ..
rapidly advancing knowledge allowed ecophysiologists
study the interactions of plant physiology and environmental
responses in the field and in laboratory growth chambers.
Animal ecophysiology developed out of. animal physiol-
ogy, concerned at first with the functioning of the human
body. V; E. Shelford (1911) stimulated the' study of animal
ecophysiology when he applied the concept of limiting fac-
tors to animals in a "law of tolerance." This law linked the
physiology of an organism to its environment. He suggested
that organisms have both a negative and an optimal response .
to environmental conditions, and that these responses influ-
ence theif distribution: This idea stimulated investigations,
notably by Kurt Schmidt-Nielson (1975), into how such
responses as thermoregulation, energy metab-
olism, and wate-r balance relate to the environmental condi-
t;ions in which animals live.
Observers noted that certain plants and animals use
chemical substances for defense, and that some plant exu-
dates inhibit the growth of associated species. They began to
investigate chemical substances in the natural world. There
were studies of the role of chemicals in species recognition,
courtShip, and defense as well as studies of the chemicals
themselves. Such work has grown into the specialized field
ofebemical ecology.
Population Ecology
As plant ecology was arising out of plant geography, other
were under way. One was the voyage of
.Charles Darwin on the Beagle, during which he collected
numerous biological specimens, made detailed notes, and
mentally framed his view of life on Earth (Figure 1.2). Dar-
win (1809-1882) observed the relationships between organ-
isms and environment. He attributed the similarities and
dissimilarities of organisms within continental land masses
and among continents to geographical barriers separating the
inhabitants. He noted from his collection of fossils how suc-
cessive groups of plants and animals, distinct yet obviously
related, replaced one another over geological time.
In developing his theory, Darwin was influenced by the
writings of Thomas Malthus (1766-1834). An economist,
Malthus (1798) advanced the principle that populations grew
in geometric fashion, doubling after some period of time.
Experiencing such rapid growth, a population would outstrip
its food supply. Ultimately the population would be restrained
by a "strong, constantly operating force-among plants and
animals the waste of seeds, sickness, and premature death.
Among mankind;'misery From this concept Darwin --
6 Part I lntroduct+o-rr --_c,_
FIGURE 1.2 Charles Darwin.
developed the idea of "the survival of the fittest" as a mecha-
nism of natural selection and evolution.
Meanwhile, unknown to Darwin, an Austrian monk,
Gregor Mendel (1822-1884), was studying in his garden the
transmission of inheritable characters from one generation of
pea plants to another. The work of Mendel would have
answered a number of Darwin's questions on the mechanisms
of inheritance and provided for his theory of natural selection
the firm base it needed. Belatedly, Darwin's theory of evolu-
tion and Mendelian genetics were combined to form the study
of evolution and adaptation, two central themes in ecology.
Two papers by G. H. Hardy (1908) and W. Weinberg (1908)
on genetic equilibrium in populations mark the beginning of
population genetics. The theoretical basis of population
genetics was advanced by Sewell Wright (1931), Sir R. S.
Fisher (1930), and J.P. Haldane (1932, 1954).
\i The Malthusian concept of population growth and limi-
- -tations stimulated the study of population dynamics. P. F. Ver-
hulst (1838) of Italy the mathematical basis for
population growth under limiting conditions. Verhulst's work,
expanded by R. Pearl and L. J. Reed (1?-29), was the basis for
the contributions of A. Lotka and V. Volterra (1926) to the
study of population growth, predation, and interspecific com-
petition. Their work established the foundations of popula-
tioP- ecology, concerned with population growth, regulation,
and intraspecific and interspecific competition. The mathe-
matical models of Lotka and Volterra were tested experi-
mentally in the Soviet Union by G. F. Gause (l934)jyjth lab-
oratory populations of protozoans, and in the United States
by Thomas Park (1954) with flour beetles. Many of the con-
cepts of population genetics have been combined with ideas
from population ecology to make up the field of evolution-
ary ecology, concerned with the interactions of population
dynamics, genetics, natural selection, and evolution.
Closely associated with population and evolutionary
ecology-and often difficult to separate from them clearly
and completely-is community ecology. More
community ecology is concerned.with interactions among
predation, parasitism, and mutual-
isui-and their influences on species abundance and distrib-
ution. More broadly community ecology also deals with the
physical and biological structure of communities, community
dynamics, and processes such as succession.
, , Ecologists now have a body of theory relating to com-
petition, population growth, life history strategies, resource
utilization, nic{hes, coevolution, community structure, fgod
webs,\and tlie like. Theoretical ecologists take theories and
equations in pure mathematics, physics, and even
economics and apply them to ecological questions. They
attempt to provide a substantial mathematical foundation for
ecological concepts, upon which predictions can be based.
Theoretical ecologists have stimulated new insights into rela-
tionships among species, utilization of resources, and life his-
tory patterns. Critics of theoretical ecology argue that it
suffers from too many hypotheses that are untested or
untestable in the field.
Ecosystem Ecology
Early ecologists, particularly plant ecologists, were con-
cerned with observing the patterns of organisms in nature,
attempting to understand how patterns were formed and
maintained by interactions with the physical environment.
Some, notably F. E: Clements, sought some system of orga-
nizing nature. He proposed that the plant community
behaves as a complex organism or superorganism that grows
and develops through stages to a mature or climax stage. His
idea was accepted and advanced by other ecologists. A few
ecologists, however, notably A. G. Tansley (1871-1955),
did not share this view (see page 8). In its place he advanced
a holistic and integrated ecological concept that combined
living organisms and their physical environment into a sys-
tem, which he called an ecosystem. A system is a complex
in which the parts interact to produce the behavior of the
whole.
Although Tansley formally advanced the concept of the
ecosystem, limnologists in both Europe and America were
using a holistic ecosystem type approach to the study of ecol-
ogy of lakes that predated Tansley. Prominent among these
scientists were A. Thienemann and F. A. Forel (1841-1912).
Thienemann developed an ecological approach to freshwater
biology. He introduced the ideas of organic nutrient cycling
and feeding levels, using the terms producers and consumers.
Forel was more interested in the physical par_3:1Il(!!ers of fresh-
water habitats, especially lakes. In his monograph on Lake
Leman, he introduced term limnology for the study of fresh-
water life.
Concepts of lake dynamics were further developed by
S. A. Forbes ( 1844- 930), an entomologist at the University
of lllinois and the lllinois State laboratory of Natural History
(illinois Natural History Survey), which he founded in 1878.
He wrote a classic paper of ecology, ''The Lake as a Micro-
cosm" (1887). It concerned the interrelations of life in a lake,
particularly through food c;hains, and the role of natural
selection in the regulation of numbers of predators and prey ..
Thus limnological approaches to the study of lakes con-
_tributed the development ofecosystem ecology.
Unrelated to limnology, but destined to have an impor-
tant influence on its future and that of ecology, was the work
of Edgar Transeau in an lllinois cornfield. Transeau was not
an ecologist, much less a limnologist. He was interested in
improving farm production by understanding the photosyn-
thetic efficiency of the corn plant. His landmark paper, ''The
Accumulation of Energy in Plants" (1926), marked the begin-
ning of the study of primary production and energy budgets.
Thienemann's and Transeau's work stimulated the study
of lakes by E. A. Birge and by C. Juday of the Wisconsin
Natural History Survey. In a classic paper, ''The Annual
Energy Bulfget of an Inland Lake" (1940), Juday summarized
the accumulation of energy by aquatic plants over a year and
also its movement through various feeding groups, including
the decomposers.
The work of Juday and Birge influenced R. A. Linde-
man, a young limnologist at the University of Minnesota.
Lindeman was interested in exploring plant succession in
terms of energy. He turned his attention to Cedar Bog Lake
in Minnesota. In a 1942 paper, "The Trophic-Dynamic
Aspect of Ecology," Lindeman described I succession in
terms of energy flow through the lake He
showed how short -term proeesses of feeding, trophic tela-
tionships, affected the long-term changes in' the lake.
paper, a significant advance in ecology, marked the
ning of ecosystem ecology.
Preceding Lindeman's contribution was the theoretical
work of a physical chemist, A. J. Lotka. In his book Elements
of Physical Biology (1925) he introduced thermodynamic prin-
ciples of energy transformations in biology along the lines of
physical chemistry. He considered food webs and the cycles of
carbon dioxide, phosphorus, nitrogen, and water; and he
viewed Earth as a single energy-transforming system. Most
ecologists overlooked Lotka' s contribution. However, his ideas
and Lindeman's study stimulated further pioneering work on
energy flow and nutrient budgets by G. E. Hutchinson (1957,
1969) and H. T. and E. P. Odum in the 1950s. J. Ovington
(1962) in England and Rodin and Bazilevic (1967) in the
Soviet Union investigated nutrient cycling in forests. The
increased ability to measure energy flows and nutrient
cycling by means of radioactive tracers and to analyze large
amounts of data with computers permitted
of systems ecology, the application of generalsystems-the""
ory and methods to ecology.
Chapter 1 Ecology: Meaning and Scope 7
Long-Term- Ecological Research
For many ecological processes, long periods of observatio:q
are required to deted:\:p.anges. For such processes long-term
research is needed. Many patterns and processes do not vary
through time in a fashion. Series of observations
must be collected for any trend or pattern to emerge.
Unfortunately long-term studies are uncommon, despii.g:
repeated evidence of the misleading nature of short-term
research. Short-term funding lies at the root of the problem.
Demands of both academic institutions and research funding
agencies for scientists to obtain results and publish papers
quickly precludes any projects running three to five years;
most are for one to two ,years. Further complicating the situ-
ation are the processes that control tenure and advancement
at academic institutions, which operate on time scales much
shorter than most ecological processes.
To address the lack of organized research efforts over
long time scales, the National Science Foundation (NFS)
initiated ,a program in Long Term Ecological Research
(LTER) ,in 1980. Researchers at a network of 21 sites across
the United States now design studies on time scales of
years, to decades, to a century (see Bioscience 1990,
40:495-523). They also address a wide range of spatial
scales-meters to kilometers to cross-continent compar-
isons among research sites.
Sites in the LTER system currently extend from Puerto
Rico to northern Alaska and represent a broad diversity of
environments and ecosystems, including temperate and trop-
ical forests, prairie, desert, alpine and arctic tundra, agricul-
.. hl!al fields, lakes, rivers, coastal wetlands, and urban sites. All
sites are large enough to incorporate moderate- to large-scale
landscape mosaics, and a majority of these site include human-
manipulated as well as natural ecosystems. In 1997, the LTER
network expanded to include two ne:r sites, Baltimore, MD,
and Phoenix, AZ. The objective of :adding urban sites is to
exrunme the interactions between ecological and socioeco-
nomic components of the :urban environment, a growing area
of concern since urbanization continues to increase.
The LTER sites are as varied in investigative design and
objectives as they are in the range of ecosystems they encom-
pass. To provide a focus for the individual projects, core
research objectives were established early in the program. All
LTER studies must at heart support these core objectives.
These objectives are to understand the patterns and controls
of primary production, food webs, population abundance and
distribution, organic matter accumulation, and biogeochem-
ical cycling as well as to answer questions related to distur-
bance frequency and effect. Research approaches include
observation, experimentation, comparative analysis, retro-
spective studies, and dynamic modeling.
Our current understanding of how key ecological
processes, such as primary productivity and decomposition,
vary among ecosystems comes laii<?lY piecing
together disparate studies. ConsequentL)4-1he-development
- of comparable data sets and standardization iiiaiHil)'ticai
methods and equipment were concerns addressed at the
::r-- Part I Introduction
beginning of the LTER program. This approach permits
comparative studies to take place within the LTER network
and to extend to non-LTER projects. Coordinated and coop-
erative research across diverse ecosystems take many
forms, including the installation of standard experimental
designs across many sites. Such integrated intersite com-
parisons are essential to addressing questions concerning,
for example, climate change and its potential influence on
key ecological processes.
TENSIONS WITHIN ECOLOGY
The complexity of ecology's past has led some scientists into
opposing camps. Often these fundamental divisions have
been, or can be made, productive.
. q
Plant Versus Animal Ecology
The first major split in ecology was the failure of plant ecol-
ogy and animal ecology to meet on common ground. In Eng-
land plant ecology was influenced strongly by A. E. Tansley
and animal ecology by Charles Elton. At that time one
nal, The Journal of Ecology, sponsored by the British Eco-
logical Society, covered the field of ecology. In a few years
Elton started The Journal of Animal Ecology. The two, plant
ecology and animal ecology, went their separate ways.
In the United States the split was less amicable. Early
on, a controversy developed over the term ecology.
Botanists decided at the Madison (Wisconsin) Botanical
Congress iri 1893 to drop the o from oecology and adopt an
anglicized spelling. Zoologists refused to recognize the
term at all. The entomologist William Morton,Wheeler
complained that botanists had usurped the and had
distorted the science. He urged zoologists to drop the term
and adopt the word ethology.
The schism was widened by a more fundamental dif-
ference in approach. Plant ecologists ignored any interac-
tion between plants and animals. In effect, they viewed
plants as growing in a world without parasitic insects and
grazing herbivores. For years plant and animal ecologists
kept to their ,separate ways. F. E. Clements and V. E.
Shelford began to bring the two !!ides together with Bio-
ecology (1939), in which they suggested that plants and ani-
mals be considered as interacting components of broad
. biotic communities or biomes.
Ji
Organismal Versus lnd,ividualislic Ecology
Although the division between plant ;md animal ecology nar-
rowed, a new division was to It had its roots
in the ideas of Clements, who strongly influenced philo-
sophical ideas in ecology. Clements viewed the plant com-
munity as an organism. Like an individual organism,
vegetation moved through several stages of development,
from youthful colonization of bare ground to a mature, self-
reproduclng climax in with its climate-determined
environment. The climax was the end or goal toward which
all vegetation progressed. If disturbed, vegetation responded
by retracing its developmental stages to the climax again.
Clements's organismal approach was not lost on animal
ecologists. In the United States the zoologist and aninial
behaviorist Williari.l Morton Wheeler, an international author-
ity on ants and termites, advanced the idea that ant colonies
behave as organisms. They carry out such functions as food
gathering, nutrition, self-defense, and reproduction. Basing
his ideas on those of C. Lloyd Morgan, a biological philoso-
pher, Wheeler applied emergence theory to ecology. He pro-
. posed that natural associations have certain emergent
properties as aggregations of organisms-predators and prey,
parasites and hosts-that arose from lower levels of organi-
zation. All levels occurred together in an ecological commu-
nity or biocenosis. The biocenosis modified its component
'> species through behavioral changes and new levels of inte-
gration. Everything in the biocenosis was related to. every-
thing else. His view of a tight but orderly nature contrasted
with'the chattic impacts imposed o_n nature by humans.
This organismic, levels-of-hierarchy view of natilre
advanced by Clements and Wheeler captured the thinking of
that influential group of ecologists at the University of
Chicago, the authors of The Principles of Animal Ecology
(1949), Allee, Park, Park, Emerson, and Schmidt. In that book
they stated that the organismic concept of ecology was "one
of the fruitful ideas contributed by biological science to mod-
em civilization."
Although the organismal concept dominated ecology
until the early 1960s, many ecologists refused to accept it.
Clement's organismic concept had its critics, notably H. A.
Gleason and A. G. Tansley. In 1926 Gleason published "The
Individualistic Concept of the Plant Association." In it he
argued that the plant association was hardly an organism
capable of self-reproduction. Instead, he argued, each com-
munity is unique. It arises randomly through environmental
selection of seeds, spores, and other reproductive parts of
plants that enter a particular area. Thus no sharp boundaries
exist between plant communities. Plants exist along a con-
tinuum dictated by changing environmental conditions. The
English ecologist A. G. Tansley, once enamored with the
organismic concept, ultimately rejected it too. Vegetation, he
allowed, might be called a quasi-organism, but certainly not
an organism or a complex organism. In fact, Tansley rejected
_the whole idea of a biotic community as anthropomorphic.
No social relationship exists among plants or between plants
and animals as the term connotes, he argued. In its place
Tansley substituted the term ecosystem. He viewed plants
and animals as components of a system that also included
physical factors.
Holism Versus Reductionism
By the mid-1960s the individualistic concept of Gleason
had supplanted the organismic concept-almost. Many of
its philosophical and functional attributes lived on in the
"new ecology" of the new ecology, as defined
by E. P. Odum (1964, 1971), is a "systems-ecology," an
discipline that deals with supraindividuallev-
els of organization."
According to this concept ecosystems develop from
youth to maturity. Each stage of development exhibits some
of its own unique characteristics. Interactions among popu-
lations and between plants and animals result in a hierarchi-
cal organization. This organization involves interacting
components that produce large functional wholes. The out-
come is the emergence of new system properties that are not
evident at the level below (Odum 1971, 1982). These emer-
gent properties account for most of the changes in species and'
growth that take place over time. The approach is holistic
(studying the total behavior or attributes of a complex sys-
tem) because systems are considered too complex to study in
bits. Because the whole is greater than the sum of its parts,
ecosystems can be studied only as functional units.
This holistic approach has critics who take a reduction-
ist approach. They consider that the ecosystem is the sum of
its parts. By understanding how each part-the species, their
numbers, and characteru,tics-functions, we can discover
how the whole system opbfates. Rather than guiding the evo-
lution of species, the nature of ecosystems results from the
evolution of species.
Fenchel (1987: 17) puts the reductionist' s point of view
well: "I find the entire argument as nonsensical as stating that
an alarm clock is qualitatively different from its constituent
wheel, bolts, and springs. A holist approach to an alarm clock
... is to observe that when wound it will run. To arrive at a
real understanding of the device one must take it apart in order
to see how it works ... to take a reductionist's approach."
The holist would counter that studying the wheels, bolts,
and springs tells nothing about the way the whole system
functions, what the clock really does. You could study a few
separate components, but they are outside of the
whole clock. Only when all parts of the systeJ\n are fuDction-
ing as a unit can the clock function. Then its prop-
erty, telling time, becomes apparent. \
Is the sum of the parts of the clock greater than the whole
or not? Allen and Starr in their book Hierarchy (1982) argue
that the whole problem of emergent properties is a matter of
scale and assert some properties of whole are
and cannot be denved from the behav10r of the parts alone.
They also point out that ecosystem models of holists are
ply large-scale reductionism. Ecosystem ecologists cannot pos-
sibly study a model of an entire ecosystem. For one, boundaries
of most ecosystems are arbitrary. Further, ecosystems are not
closed. One ecosystem feeds into another. Thus ecologists can
only study pieces of it. The only major difference between a
reductionist and a holist is that the holist studies larger pieces,
made up of assembled parts studied by the reductionists.
What keeps ecosystem ecologists (holists) and popula-
tion and evolutionary ecologists (reductionists) apart is their
approach to ecology. Population ecologists focus on species'
interactions_with their environment in the broadest sense.
They are-interestedin the historlcal-o-r-ultimatereasons why
natural selection favored different adaptive responses among
species over evolutionary time. Ecosystem ecologists are
Chapter 1 Ecology: Meaning and Scope
more interested in the how of current or proximate outcomes
of the functional interactions at the population, community,
and ecosystem These differences in approach may not
be as great as they appear.
What' can bring the two groups together? Population
ecologists could approach population growth and population
interactions such as muiualism, parasitism, predation, '
competition as interacting systems (Berry 1981) and as com:::
ponents of a hierarchy of systems. Systems ecologists could
integrate some evolutionary theory into system models, par-
ticularly in the area of ecosystem development and organiza-
tion (Loehle and Peckmann 1988). Food .Web theory, for
example, crosses the line into both evolutionary and systems
ecology, involving botlt species interactions and the transfer
of energy and nutrients through a hierarchy. Ecosystem func-
tioning ultimately depends on specil::s adaptations, which are
the outcomes of evolution. For example, efficiency of water _
use by certain ecosystems such as grasslands and deserts
results from the water use efficiency of the individual plants.
The natu:tal assemblage of plants and animals that make up
the living component of an ecosystem is not a random col-
lection of species; rather it is one that has been determined
by the competitive abilities and other attributes of the com-
ponent species (H. Odum 1983).
APPLIED ECOLOGY
Ecological theories and models help us understand the
human impact on environments. They provide a basis for
... ecosystem and natural resource management, preservation,
. and restoration. All these activities make up applied ecol-
ogy. For years theoretical and academic ecologists viewed
applied ecology as an intellectual lightweight. Applied ecol-
ogists, for their part, often ignored theory, even when it
be of practical use; Fortunately ecologists of both per-
now recognize that solutiqns applied to environ-
mental problems must be based on sound theory developed
through research.
Applied ecology began to take shape in the 1930s. In
1932 Herbert Stoddard pointed out the role of fire in the con-
trol of plant succession in his book The Bobwhite Quail. This
topic was ignored by academic plant ecologists. Aldo
Leopold pioneered the application of ecological principles to
the management of wildlife in his classic Game Management
(1933). In Forest Soils (1954), H. L. Lutz and R. F. Chandler
discussed nutrient cycles and their role in the forest ecosys-
tem. J. Kittredge pointed out the impact of forests on the envi-
ronment in Forest Influences (1948).
Although applied ecology has been around since the
early 1930s, it did not gain visibility until the 1970s, when
ecology became involved in social, political, and economic
issues. This_i_nvolvement grew out of public awareness of the
problems ofpolliition, toxic wastes, overpopulation, and a
degraded-environm.e_nt. Although-the- public treated these
issues as if they were new,-ecOIOgists had grappled with envi-
ronmental problems for years. The ecological movement had
10 Part I lntrodm:ti!in
its roots in Europe, especially Germany. An early founder of
political ecology was Ernst Haeckel. From Germany it moved
to northern Europe, Great Britain, and the United States. In
England the animal ecologist Charles Elton helped found the
Nature Conservancy. The plant ecologist A. G. Tansley
founded the British Ecological Society and was active in the
conservation movement.
In the United States George Perkin Marsh called attention
to the effects of poor land use on the human environment in
his dramatic book Man and Nature (1885). In the 1930s F. E.
Clements urged that the Great Plains be managed as grazing
land and not be broken by the plow. The plant ecologist Paul
Sears wrote Deserts on the March (1935) in response to the
Great Plains Dust Bowl of the 1930s. William Vogt's Road to
Survival (1948) and Fairfield Osborn's Our Plundered Planet
(1948) called attention to the growing population-resource
problem. Aldo Leopo,d's A Sand County Almanac (194;9),
which called for an ecological land ethic, was read largely by
those interested in wildlife management until the 1970s, when
it became the bible of the environmental movement.
Rachel Carson did more than anyone else to bring
ronmental problems to the attention of the public (Figure- L3).
Since the publication of her book Silent Spring (1962), peo-
ple have become more aware that chemical poisons and other
FIGURE 1.3 Rachel Carson.
pollutants are recycled through the environment. Once casti-
gated as more fiction than fact, Carson's predictions came
only too true as carnivorous birds fell victim to toxic chemi-
cals. With a ban on DDT in the United States, some eagles,
hawks, and osprey began a gradual comeback. Carson made
people quick to recognize other continuing chemical dangers,
such as dioxin and PCBs.
The alarming decline in species populations, the threat-
ened extinction of many forms of life, the fragmentation and
.loss of habitat, and the burgeoning impact of human popula-
tion growth on Earth's natural resources have impelled oth-
erWise academic ecologists into applied ecology to become
involved in resource management. Their entrance into applied
ecology has resulted in a growing interest in and effort to
apply ecological theory to environmental problems. This
is reflected in the development of three new fields of
biology, restoration ecology, and
landscape the establishment of new journals in
the 'field-qonservation Biology, Landscape Ecology,
Restoration Jf,cology, Journal of Applied Ecology, Ecological
ApplicationS, Environmental Management, and Ecosystems.
Conservation biology is a new synthetic field that
applies principles of many disparate fields-ecology, bio-
geography, population genetics, economics, sociology,
anthropology, philosophy, and other fields-to the mainte-
nance of biological diversity (Meffe and Carroll 1997). It
aims to generate new scientific approaches by melding pop-
ulation biology with applied fields of ecology to solve the
problems of protection and maintenance of biodiversity in the
face of homogenizing and destructive forces affecting many
ecosystems. Although conservation biology in some ways
stands apart from other applied fields, its ultimate success
will require its joining in part with. forestry, range manage-
ment, wildlife and fishery management, and resource eco-
nomics. In some ways it has its roots in wildlife biology, a
field in which biological knowledge and ecological principles
were first applied systematically to conservation of organisms
and their natural habitats.
Restoration ecology involves the application of prin-
ciples of ecosystem development and function to the
restoration and mapagement of disturbed lands (see Jordan
et al. 1987, Baldwin et al. 1994, Hobbs and Norton 1996,
Meffe and Carroll 1997). Its goal is to return a particular
habitat or ecosystem to conditions as similar as possible to
the predegraded state. Restoration ecology covers a con-
tinuum from the reclamation of highly disturbed local sites
and the re-creation of a particular ecosystem to restoration
and maintenance of entire landscapes for sustainable pro-
duction and conservation values. The restoration of
degraded land means rebuilding functional ecosystems, but
not necessarily restoring the site to resemble the original in
all its aspects. Restoration ecology offers ecolo-
gists the opportunity to learn about ecosystem structure and
function by putting them back together, to develop guiding
principles for restoration, a{_ld to test ecological ideas.
Learning what-does and ".'hat does not work provides
insights into how ecosystems function. Although restora-
Chapter 1 Ecology: Meaning and Scope 11
tion ecology has been concerned primarily with small on- degradation, including the loss of species. However
site reclamation, restoration ecologists are moving in the most definitions envision ecosystem management as requir-
direction of a much larger scale-that of the landscape. ing the integration of ecological, socioeconomic, and insti-
Restoration ecology is also concerned with reestablishing tutional perspectives.
landscape patterns and heterogeneity that have been altered Although a of ecosystem management are
by disturbance and increasing conservation values in frag- being implemented by 'some governmental agencies on pub-
mented or modified' landscapes. This expanded interest is licly owned lands and by some nongovernmental institutions,
bringing restoration ecology into a close relationship with ecosystem management:has its vocal and often bitter critics.
landscape ecology. At the heart of the controversy is the fear among developer!!i
Landscape ecology is the ecology of landscapes-land many foresters, range managers, and others that ecosystem
areas composed of clusters of local ecosystems repeated in a management with its emphasis on maintaining full array of
similar manner across kilometers-wide areas, such as ecosystem functions threatens short-term economic gains.
of forest existing in an agricultural or urbanized landscape. They seeit as marking the end of traditional, commodity-on-
Thus landscape ecology is concerned with spatial patterns in ented monocultural use of land, threatening takeover of the
the landscape and how they develop,/ with an emphasis on the management of private forests and ranges, and placing con-
role of disturbance, including human impacts (Fo111;lan and trois on development. Critics have cultivated in the public a
Godron 1986, Forman 1995, Turner 1989, Farina 1998). It mistaken belief that ecosystem management is a program
explores how a heterogeneous combination of ecosystems is designed as a way for federal government to expand
structured, how it functions and changes over time. It studies ity over state and local land use. This, of course, is wrong .
the distribution of landscape elements, such as fields, forest, because federal and state agencies require input from the pub-
grassland, highway rights of way, and suburban subdivisions, lie, and management of resources falls under the aegis of fed-
and their spatial and temporal interactions and exchanges eral, state, and many local governmental agencies. For good
across ecosystem boundaries-movement and flows of plants, of ecosystem management, see Grumbine (1994)
animals, energy, mineral nutrients, and water. and Meffe and Cannll (1997).
Beginning as a somewhat undefined subunit of ecol- , . In the past quarter century since people became con-
ogy, landscape ecology has expanded greatly. Few ecolo- cerned about growing environmental degradation, how has
gists would undertake a community or ecosystem study the situation changed? We started off well enough with envi-
without considering landscape-level implications. It has ronmentallegislation: the National Environmental Policy Act
become an integral part of forestry, wildlife management, (1969), designed to protect the environment from overzeal-
ecosystem management, and conservation biology, all deal- ous development and to mitigate losses, the Endangered
ing with landscape-level interactions. Landscape ecology Species Act (1973, amended 1982 and 1994), the Clean
employs the recently developed techniques of geographic Water Act (1977, amended 1981, 1987, 1994), and the Clean
information systems (GIS). These are computerized sys- AfrAct of 1970 and 1977, among others. The early enthusi-
tems that generate maps containing spatial_ information asm for a quality environment is still strong with the public.
about a portion of a landscape, such as topography, vege- It has weakened, however, among many politicians; govem-
tation patterns, and other physical features tbat that-tan be ment is less sensitive about environmental issues. During the
displayed from remote sensing data. The information is 1980s there was even an environmeiltal backlash at the fed-
stored digitally and can be presented or graphi- erM level, as the Reagan administration attempted but failed
cally, making it available for comparing various landscape to undo all the environmental progress made during the pre-
features at different locations or at the same location over vious two decades.
time. Such maps can be linked to population simulation There has, of course, been progress. Water quality has
models that enable ecologists to predict potential responses improved considerably, and the air above some of our cities
of different organisms to specific landscape changes is cleaner. However, we have discovered that our environ-
Ham et al. 1992). mental problems are not only more difficult to solve than
We have begun to apply ecosystem and theoretical once believed; many are growing worse. Toxic wastes pol-
ecology more intensively to resource management in the lute groundwater and land. Air is becoming more polluted
past decade, even though economics too often takes prece- worldwide. Haze has cut visibility in the eastern United
dence over sustainability. A recent approach is ecosystem States by more than 50 percent in the past 40 years. Acid
management, grounded in landscape ecology. Like ecol- rain affects lakes and streams. Increased concentrations of
ogy, the term ecosystem management has a diversity of def- carbon dioxide and ozone threaten climatic stability. Roads
initions. A simple but perhaps inadequate definition is the cut into open country, and suburban expansions eat away at
following: Ecosystem management considers ecological the hinterlands and farmlands. Continued deforestation in
systems as functional units and stresses their long-term sus- both temperate and tropical regions is fragmenting wildlife
It emphasizes a shift from short-term yields and habitat, increasing the rate of extinction. A rapidly growing
economic gains to long-term management that sustains or .urban and suburban population with increasing interest in
_____ restores a natural O_!}nodifieu ecosystem at a level . outdoor recreation intolerable pressures on
allews-human use but does not result in-long-term ecosystem and'-nationaLparks that threaten-their eeological integrity.
------------------ -- - -
12 ParH lnt!'oduction
FIGURE 1.4 Human activities
lessen the sustainability of the
biosphere.
Even the oceans have not escaped, as human debris and
chemicals have been deadening the seas and destroying
marine life. In spite of surplus agricultural production, wet-
lands are still being drained for more cropland at an alarming
rate, threatening the existence of already dangerously
declining wetland wildlife. All these activities have affected
regional and global ecological processes and have lessened
Earth's ability to support a diversity of life, including
. humans (Figure 1.4). With the human population growing
at the rate of 1.8 percent annually, its pressures on Earth's
resources will accelerate.
Among the many environmental problems facing
humanity, four broad areas are critical: global climate change,
biological diversity, sustainability, and rapidly growing
human populations. We are causing dramatic changes in the
distribution, abundance, and number of species. The loss of.
diversity can affect the stability of communities and popula-
tions on which our economy depends, as exemplified by the
loss of commercially important fish species. This rapid
climinution of E!)rth's resources affects our ability to sustain
both natural and managed ecosystems and human life itself.
The basis and solution of our environmental problems
are ecological in nature. To this end The Ecological Society
of America has developed a three-pronged Sustainable Bios-
phere Initiative, involving research, education, and environ- .
mental decision making (Lubchenco et al. 1991). Research
priorities focus on the critical areas of global change, bio-
logical diversity, and sustainable ecological systems.
Researchers seek answers to such problems as the responses
of ecological systems to stress, development and application
of ecological theory to the management of ecological sys-
. terns, and an ecological understanding of the effects of intro-
Sttuced species, pests, and pathogens.
Unfortunately, attempts to apply sound ecological prin-
ciples to environmental problems often run headlong into
economic, political, and social opppsition, as witnessed by
the debates over old-growth forests;\'regulation of fishing,
land zoning, and wetlands preservation. Successful applica-
tion requires a citizenry that understands .ecology and its
importance. We need ecological education at all levels. Eco-
logical principles need to be clearly understood by econo-
mists, engineers, lawyers, businesspeople, and politicians, all
of.themdecision makers who can hurt or improve the envi-
ronment. Most decision makers are unaware of the facts, do
not understand basic ecological concepts, or are even hostile
- 'e
to environmental considerations for political, economic, or
special reasons. An educated public can cause decision mak-
to be more responsible.
The of human life on Earth depends on far more
ecological knowledge than we now possess, even though we
are n!M: all we know. For the first time in the history
of Earth, Homo sapiens has become the completely dominant
organism, changing Earth and its diversity of life at will with
little regard for the consequences. It is little wonder, then,
that some of the most intellectually challenging problems in
ecology lie in that transition zone between theoretical and
applied ecology.
ECOLOGY: AN EMPIRICAL
AND EXPERIMENTAL SCIENCE
Early on, ecology was largely observational and descriptive.
In North America so little was known about the nature and
species composition of natural communities that ecologists
concentrated on describing them and collecting factual infor-
mation. They had little incentive or basis to undertake exper-
imental studies. In Europe most ecologists concentrated on
vegetation classification. Early steps at empiricism in North
American ecology began with F. E. Clements. Studying
development of vegetation in the midwestern grasslands,
Clements recognized the inadequacy of simply observing
vegetational changes. If he was to make progress in the study
of vegetation dynamics, he had to devise some means of
quantifying and recording changes over time and the differ-
ences between various areas. To do so, Clements devised the
quadrat method (see Appendix A), which is still the sampling
unit for many ecological studies today (Tobey 1981). His
quadrats were square plots of varying sizes in which all plants
could be listed, counted, and even mapped on grids within the
quadrats. The quadrat method provided a distinct advantage
in study of vegetation .. The establishment of permanent plots
allowed the study of the development of vegetation or suc-
cession on denuded plots, the changes in plants in response
to environmental changes, and the changes in plant commu-
nities across a gradient. In spite of his quantitative approach
to the study of vegetational Clements had little
timdor..statistics, which he cobsidered as simply expressing
known facts without uncovering anything new. And, unlike
A. E. Tansley, he opposed mathematical models in ecology.
A small group of ecologists in England and
United States, however, were developing mathematical mOdels
based on theLotka-Volterraequations (see Chapters 10, 13, and
14) to study population growth, predator-prey relationships, and
competition. Much of the impetus for ma!hematical models was
directed toward applied ecology: pest control and fisheries.
Empirical methods came into own in-the 1960s. Faced
with the rapid emergence of molecular biology, the growth of
physics, and the expressed opinions that ecology was a soft sci-
> .
ence,some ecologists-notably George E. Hutchinson, Robert
MacArthur, Richard Levins, and Robert May-attempted to
revolutionize and transform ecology into a highly predictive sci.,,.
ence with its own fundamental laws like those of physics (see'
Kingsland 1995). The result was a flurry of building mathe-
matical models, many of them little more than mathematical
exercises. Ecologists ignored the fact that they were dealing not
with objects as in physics but living organisms, highly variable
in their physiology, genetics, evolutionary history, and envi-
ronmental interactions. This variability precluded ecology from
becoming an exact science. After several decades of divisive-
ness much of the furor has died. It did bring to ecology, how-
ever, a strong emphasis and hyJ>?thesis
testing, and on development of models that would provide
insights into and suggest solutions to ecological problems.
The specific goal of ecology is to understand the patterns
and processes related to life on Earth-how the variety of plants
and animals have adapted to the various environments and how
they interact among themselves and with their abiotic environ-
ment To gain some insight into these observed processes and
patterns, ecologists, like all scientists, develop possible expla-
nations about the causes of observed phenomena These possi-
ble explanations are called hypotheses. A hypothesis is a .
statement about an observation that can be tested.
There are two approaches to the development and test-
ing of hypotheses: inductive and deductive. In the inductive
method, the scientist gathers empirical and f{.pm it
arrives at a generalization. The inductive me.thod proceeds
l
from specific observations to a general conclhsion. Using a
deductive method, a scientist develops a general idea
a phenomenon, performs experiments, and from them makes
specific predictions that can be _tested again. The experi-
menters go from a general idea to a specific prediction.
Although the distinction between the two 11?-ay
seem. subtle, they are fundamentally different approa.ches.
The inductive approach is the formation of general principles'
from specific observations; the deductive approach is the pre-
diction of specific events from general principles. Each
method has its advantages, and ecologists employ both.
Experimental Approach
Testing hypotheses entails first the collection of data by
observation or experimentation. Ecologists employ direct obser-
vation and comparison in "natural experiments" (Diamond
1986). For exl.!Illple, ecologists may be interested in studying
the relationship between the abundance of standing dead trees
(snags) and the population de11sity of woodpeckers in various
forest stands. Because woodpeckers use snags as nest sites, the
Chapter 1 Ecology: Meaning and 13
hypothesis is the abundance of snags represents a limiting
resource to population size. By censusing the woodpecker pop-
ulations and the number, of dead trees in various forest stands,
the ecologist could compare the woodpecker population with
the number of snags. woodpecker density is higher on sites
with more snags, then the observations support the hypothesis.
The observed patterns, however, only suggest a correla-
tion (co-related); they do (lot address the question of cause and
effect. There may be other reasons for the variation in .
peeker population density, which may also be related to
density of snags. If the density of large trees on which to feed
limits the;population, you might expect a relationship between
. woodpecker population density and stand age. In this case, the
factor causing the variation in population density among stands
is the availability of suitable foraging sites and>not nesting sites.
It just so happens that the. abundance of nesting sites (standing
dead trees) is correlated to the causal variable of food abun-
dance (stand age). To help determine cause and effect, an
ogist may employ an experimental approach in which one or .
more variables are directly manipulated. This rigorous estab-
lishment <;>f causation is what separates science from unwar-
ranted assUm.ptions about reasons for observed phenomena
. The' experimental approach, unlike the collection of
observations from unmanipulated systems, directly deter-
mines the response of one variable-the dependent vari-
able-to variation in some other variable(s)-independent
variable(s). The investigators control the independent vari-
able. They manipulate the variable in a predetermined way,
called a treatment, and monitor the response of the depen-
dent variable. For example, in a laboratory experiment to test
a plant's growth response to increased C0
2
, the concentration
of C0
2
is the independent variable and plant growth is the
dependent variable (Figure 1.5).
0.30
...

rn
gj

:0 0,.25
Ol
.5
=a
Q)
Q)
rn
]j
t2
0.20
350 420 490 560 630 700
C02 concentration (ppm)
, FIGURE l .. ThE:! effect of atmospheric C0
2
concentration on the
growth of birch seedlings in the greenhouse. Growth is
measured as the accumulated biomass over the period of the
experiment. Each point-represents-the biomass of a seedling grown
under the corresponding atmospheric C0
2
concentration.
Part I Introduction
One important constraint in experiments is the need to
control the variation in other independent variables that may
influence the response of the dependent variable. For exam-
ple, in an experiment testing the response of plant growth to
elevated C0
2
, the investigator must be sure that all other fac-
tors, such as moisture availability, temperature, and available
light, that may influence plant growth are the same for all
treatments. Otherwise the researcher cannot definitively inter-
pret the plant response as a direct function of C0
2
alone.
Even though the investigator tries to reduce all possible
differences among individuals or units used in the experiment
and to maintain a rigid control over environmental conditions,
there will always be some amount of variation among indi-
vidual plants. One source of variation very difficult to con-
trol is the inherent genetic variation among individual
organisms in a population. Because genetic and many other
variations often cannot be controlled, the investigator must
examine the response 9f a number of individuals rather tlian
relying on a single observation. This approach is known as
replication; the individuals receiving the treatment are the
replicates. The investigator must determine the number of
replicates required for each experiment to account for_ this
uncontrolled variation. The decision will be related to the
degree of variation exhibited by individuals within the popu-
lation being examined. ,
Because the purpose of the experiment is to examine the
response of the individuals or experimental units to the treat-
ment (variation in the dependent variable), a group of indi-
viduals must be used as a control. The control is a group of
individuals (replicates) that do not receive the treatment, but
otherwise are handled exactly like the treated plant. The con-
trol forms the basis for comparison with the individuals
receiving the treatment. In the case of the experiment to
examine the response of plants to elevated C0
2
, the investi-
gator would grow the control individuals under current nor-
mal (ambient) concentration of C0
2
, thereby providing an
estimate of (expected) growth for individuals in the popula-
tion under baseline conditions.
Field Experiments
Ecologists conduct experiments in the laboratory and in the field
(Figure 1.6). Laboratory experiments usually involve synthetic
communities of one or a few species. On the basis of these sim-
plified systems, such experiments -can reveala wide range of
outcomes or relationships that the investigator can eval-
uate in the field under more natural or complex conditions.
Field experiments involve the manipulation of one or
more independent variables in a natural system. The investi-.
gator accomplishes this manipulation by adding or removing
\a species, by erecting exclosures to prevent access to a space
or resource, Of by adding or withholding essential resources.
In, contrast to laboratory experiment, the investigator often
finds\it imposfible to control the many independent variables
that may influence the response of the dependent variable.
This circumstance limits the investigator's ability to assign a
causal relationship between the response of the dependent
variable and the treatments. To improve this situation, the
investigator must include a set of experimental units or plots
as controls. These control units must be similar to those
receiving the treatment. The investigator uses this set of field
controls for comparison with the set of units on which the
treatments are applied. This comparison will form the basis
for testing the hypothesis regarding the influence of the inde-
pendent variable on the dependent variable.
Field experiments are generally on a larger scale than
laboratory experiments, and in some ways they are more
"realistic" in that the objects mi.tler study have not been iso-
lated and controlled in a laboratory environment. However,
there are limitations to field studies. One such limitation is
FIGURE 1.6 Although ecologists carry on experimental studies in the laboratory, much of their
work is done in the field. (a) A grid established in an oak forest to study the effects of thinning and
insect defoliation on the growth and spread of Armillaria mel/ea. The fungus Armillaria me/lea
(honey mushroom) is the cause of root rot in oaks, resulting in the death of trees, especially those
of low vigor. (b) of fungal samples from soil blocks removed along the grids.
(a) (b)
the problem of adequately choosing the sample or replicates
of the objects under study. Experiments require the selection
of experimental units (such as plots or individuals), some of
which will receive the treatment (i.e., manipulation of the
independent variable) and some of which will act as the con-
trol (no manipulation). The control then serves to monitor the
natural variation in the dependent variable.
Hypothesis TesUng
Assiune the investigator has developed a hypothesis that ele-
vated levels of C0
2
result in increased-plant growth.
growth response to elevated C0
2
is the dependent variable'
and the concentration of C0
2
is the independent variable. The
investigator wants to know the effect of varying the indepen-
dent variable on the dependent variable. The next step is to
collect the data necessary to test the hypothesis and reach a
conclusion. The experimental data would consist of growth
measures (e.g., biomass gain over a period of time) for plants
grown under ambient (the control samples) and elevated lev-
els of C0
2
(treatment samples).
We test the statistically by creating a null
hypothesis (flo) and an alternative hypqthesis (Hi). The null
hypothesis is the statement of no difference between
and treatment units-that is, the independent variable has no
significant effect on the dependent variable. The alternative
hypothesis is a statement of significant difference between
control and treatment units. In the present experiment the
null hypothesis would state that the elevated levels of C0
2
have no effect on growth, because generally in an experi-
ment it is easier to disprove a null hypothesis than to prove
a hypothesis. If the results of these experiments showed no
difference in growth between the control and the treatment,
the investigator would fail to reject the null hypothesis. If
there was a difference, the investigator would not accept the
null hypothesis. The testing of the nu!l
statistical procedures, some of wh1ch are i m
Appendix A. \
Models and Predictions
After hypothesis testing, we can formulate a relationship
between a dependent and independent variable (for exam-
ple, plant growth and the level of C0
2
) to develop a model
of how this two-variable system functions. A model is an
abstract representation of the real system. It is an explicit set
of assumptions, typically formulated in a mathematical way.
When we construct and test models (examine their predic-
tions), we are in fact testing the underlying assumptions on
which they are structured. Failure of a model may well pro-
vide as much insight as its success by contributing infor-
mation about the validity of the assumptions on which the
model is based.
We can view hypotheses as models. The hypothesis that
Chapter 1 Ecology: Meaning and Scope - 15
other independent variables such as temperature. Thifmooel
is qualitative it only a of variation
(increased growth) in the dependent vanable rather than the
amount of variation. In most cases, once a qualitative rela-
tionship has been a quantitative model is sought.
The new model can be statistical or nonstatistical.
Statistical Models Statistical models are mathematical .
descriptions of data. They predict the value of the dependent'
variable based on mathematical functions. An example of a
statistical model is the simple linear regression model:
y = a + bx, where y is the dependent variable, xis the inde-
pendent variable, a is the parameter describing the y inter-
cept, and b is the parameter describing the of the line.
They intercept is the va}ue of y when xis equal to zero. The
slope of the line quantitatively relates the change in y per unit
change in x. The parameters, a and b, are solved for mathe-
matically based on the paired observations of x and y.
For example, a long-standing theory is that a close rela-
tionship exists between the density of small trees and shrubs
and the density of snowshoe hares (Lepus americana). Litvi-
atis et al.(1985) established two 49-hectare (ha) study sites in
spruce-fir and hardwood stands. They determined
snowshoe hare densities in spring and fall by mark capture-
recapture methods (see Appendix A), quantified habitat use
with fecal pellet counts, measured intensity of twig-clipping
by hares, and sampled the density of understory vegetation.
The results of their study, summarized in Figure 1.7, showed
a linear relationship between hare density and stem density.
The hares increased as understory stem density increased. The
equation (y = 0.000046x - 1.06) is a statistical
model for predicting the density of hares (y) given a value of
stem density for the plot (x). .
One important constraint of such statistical models,
because they are a mathematical description of data, is their
limited value when extrapolated beyo9d the bounds of obser-
vations. The model can be used legitimately to predict hare
derlsity for plots that have a stem density of 20 to 60 x 10
3
stems per hectare. Because no observations were made and
included beyond these bounds, we have no information as to
how hare density might vary with increasing density of
2.0 .--------------------,
1.5
J

Ql 1.0

I
0.5
y = 0.000046x- 1.06
r = 0.94
p < 0.001
plant growth will increase under elevated C0
2
is an explicit 20 30 40
based on the physiological understanding of pho- Stem per ha
50 60 X 10
3
tosyntlresis. We can test the model experimentally and either ----FIG ORE 1. 7 Relationship between stem density and.
-------accept:0r-F@jectiLFurther, we can test the generality of this snowshoe hare- dffiiSityin Maine, 1981-1983. (From L1tva1t1s
model with other species, or examine the interaction with et al. 1985.)
16 I Introduction
6000
5000
co
<E.
Ol
4000
6
c
0
n
:::l
""0
3000
e
0.
"'
"'

(.') 2000
1000
100 . 200 300 400 500 epo
1 Rainfall (mm)
FIGURE 1.8 Production of the grassland of Namibia in relation to
annual rainfall. (From H. Walter 1973.)
stems. With increasing density of stems, hare density may con-
tinue to increase, level off because some other factor now lim-
its population density, or even decline because the stem density
impedes mobility. To predict the effect of stem densities beyond
the bounds of these observations, additional plots with stem
densities above 50 and below 20 would have to be included ip.
the analysis. Another way of looking at it is that the model (the
regression equation) is a description of the correlation between
the two variables, not a statement of causation.
Nonstatistical Models In contrast to statistical models,
nonstatistical models are much less well defined. In a way
they make up an aggregate of all other mathematical mod-
. els. The main difference between these two categories is
that many nonstatistical mathematical models go beyond a
description of the relationship between the dependent and
independent viuiables and assign a mechanism to the para-
meters. For example, ecologists have used data on
land productivity from arid and semiarid grasslands to
develop a statistical model relating grassland productivity
to annual rainfall (Figure 1.8) (H. Walter 1973). Although
the model provides a good estimate ofexpected productiv- .
'i.ty (dependent variable) within the range of rainfall (inde-
pendent variable) examined, it;. does not address the
mechanism by 'Yhich this comes to exist.
To accomplish this goal, we rieed a much more com-
plex model of plant processes. An is a model for
the grasslands of the Serengeti in East Africa (Coughenour
et al. 1985). This model simulates the of photo-
synthesis and respiration as well as the cycling of nutrients
through the ecosystem (Figure 1.9). The model is made up
of a of mathematical equations that describe the
processes of C0
2
uptake, respiration, and los!;_9f Wll,ter from
leaves and other plant functions. It can predict patterns-of
net primary productivity, evapotranspiration, and nutrient
cycling through grassland ecosystems as a function of soil,
rainfall, and herbivory. Although the model addresses the
specific mechanisms responsible for the relationship
between rainfall and grassland productivity, it requires
much data to define the parameters relating each of these
processes to plant growth. Care must be taken that the
model can be-understood and interpreted. Although this
approach is a valuable contribution to understanding the
grassland ecosystem, the statistical model may, within the
bounds of observation, prove to be a better predictor for the
purposes of management. Both types of models are essen-
tiat in the development and application of science.
Analytical Models Within the large category of nonsta-
tistical models are two contrasting types: analytical and
Analytical models can be solved mathemati-
ca1ly. The inyestigator can analyze the behavior of the
model with to the parameters. Because only one
solutit>n existf for a given set of parameters, the model is
deterministic. An example is the logistic population model
(see Chapter 9)
dN!dt = rN(K - N)IK
where N is some measure of the population size such as the
number of individuals or biomass, tis' time, ris the instanta-
neous growth rate expressed in per capita units of N, and K
is the carrying capacity or the maximum sustainable value of
N for the given environment. The logistic model is analytical
because the equation can be solved. For example, consider a
hypothetical population of an initial size of 30 with the para-
meters of r = 0.186 and K =50. If we insert these parame-
ters into the logistic model and solve, we arrive at a predicted
population size of 38 in 6 years aqd 49 in 20 years. Thus the
behavior of the model can be analyzed with respect to the
parameters. Only one solution exists for a given set of para-
meters (rand K) arid initial population size (N).
Simulation Models In contrast, simulation models cannot
be solVed analytically. An example is the group of models
known as individual-based population models. In contrast to the
logistic population growth model, individual-based population
models simulate each individual organism within the popula-
, :tion rather than an aggregated parameter describing the popula-
. tion as a whole. For exlltnple, the parameter r in the logistic
equation is a popuhttidil estimate of the average net fecundity
rate per individual and -represents the carrying capacity of the
habitat in terms of maXim:illll sustainable population size. Indi-
vidual-based models, however, simul.ate the establishment,
growth, reproduction, and mortality of each individual in the
population, with these processes responding directly to avail-
ability of resources like water, light, and nutrients. Fecundity
rates may vary among individuals based on their size and age,
and the carrying capacity of the habitat may vary as a function
of resource availability. The population response is viewed as
the composite of the individuals. One advantage of this approach
is that it is not necessary to make two implicit as11!!1I:Jtions asso-
- eiated with the population-level approach typified by the logis:
tic model: (1) all individuals within the population are
Chapter 1 Ecology: Meaning and Scope 17
Grazing
FIGURE 1.9 Nonstatistica:l model of the Serengeti grassland ecosystem. The diagram depicts the
components of the system-son water, plant water, leaf area, and root biomass (designated by
rectangles )-and the processes, flows of water, and nutrient among the All
components are affected by grazing, which reduces the leaf area of and removes the growing
points at the tips of roots and stems. The squares c. and c; are respectively and internal
conductances of water. Doors indicate flow regulation. (Adapted from Coughenour et al. 1985.)
l
identical-the unique features of each are unimportant; (2) the
population is perfectly mixed-there are no local spatial inter-
actions of any important magnitude (Huston et al. 1988).
An example of individual-based models is the forest gap
model (Botkin et al. 1971. Shugart and West 1977)._ Forest
gap models simulate the establishment, growth, reproduc-
tion, and mortality of individual trees on a forrst stand. Each
individual responds to the environment of tl)e forest\stand
(temperature and available light, water, and and in
return influences the environment of the stand (for instancet
taller trees shade smaller These models are much'
more complex than the population-level models in that each
individual in the population must be simulated through time,
requiring the use of complex computer programs. The mod-
els require a great amount of information on the life histbry
of species to relate the processes of growth, reproduction,
and mortality to environmental conditions. In fact the mod-
els must simulate (1) the environmental conditions of the for-
est stand; (2) the response of plants to that environment; and
(3) how plants modify the environment of the plot, influ-
encing the other individuals in the stand.
Both modeling approaches have their advantages and dis-
advantages. The more appropriate approach depends on the
objectives of the investigator and the question being addressed.
Validation
Once a model has been developed, it must
Valioatiun is the test of how much confidence can be
placed in the model. Does the model really agree with the
behavior of the real-life system it is to mimic? Can it pro-
duce empirically correct predictions? To validate the
model, the investigators must collect an independent set of
data in the field, incorporate it into the model, and deter-
mine how well the predictions of the model based on the
new set of data compare with data used to frame the model
(Jeffers 1988). This step is important; ecology has many
models, but few are validated.
..
Summary
1. Ecology is difficult to define precisely. A simple, but not inclu-
sive, definition is the study of the interrelations of organisms
with their physical and biological environments.
2. Ecology's origins are diverse, but a main root goes back to
early natural history and plant geography. Those evolved into
the study of plant communities. European plant ecologists con-
centrated on describing plant communities. American plant
ecologists concentrated on studying the development and
dynamics of plant communities.
3. Animal ecology developed later than plant ecology. It evolved
into the study of natural selection and evolution, beginning
with the major contribution of Darwin. It gave rise to behav-
ioral ecology, which is concerned with the way animals inter-
act with their living and nonliving environment as influenced
by natural selection.
4. Studies of physiological responses of animals and plants to
temperature, moisture, light, nutrients, and other environmen- _
tal factors grew into physiological ecology. Studies-of-certain---
" cEeiiiicat-reactions of organisms to their environment stimu- -
lated the development of chemical ecology. Chemical ecology
18 Part I tntrtTIIuction
.:.-

is the study of the uses of chemicals by plants and animals as
attractants, repellents, and defensive mechanisms, and of their
evolution and chemical structure.
5. Darwin's theory of natural selection and Mendel's work in
genetics gave rise to the field of population genetics. The
Malthusian concept of popul_ation growth and limitations gave
birth to population ecology, concerned with population growth,
regulation, competition, and predation. Concepts of population
genetics and population ecology combined to form evolution-
ary ecology and theoretical ecology concerned with interac-
tions of population, genetics, natural selection, and evolution.
Closely allied with population ecology is community ecology,
which in part deals with population interactions that affect com-
munity structure and in part with the physical structure and the
dynamics and processes associated with community change.
6. As its various disciplines expand, ecology is becoming frag-
mented into specialties, often with a deepening lack of com-
munication among them. Two major divisions are holistic
ecosystem ecology, with emphasis on emergent properties/land
reductionist evolutiqnary and population ecology, which views
ecosystems as the sum of parts that can be studied separately
to discover ecosystem functions.
7. Applied ecology is concerned with the application of
cal principles to major environmental and resource manage-
ment problems. Traditionally, applied ecology meant forest;
range, wildlife, and fishery management. Recently, applied
ecology has spawned the new fields of conservation biology,
restoration ecology, and landscape ecology. The future of the
quality of human life in all its aspects and the sustainability of
Earth depend on our ability to recognize and apply ecological
principles to our use of natural resources. One approach is
ecosystem management that attempts to integrate ecological,
socioeconomic, and institutional perspectives to allow use m a
manner that will not.result in long-term ecosystem degradation.
8. Ecology has evolved from a descriptive to an empirical and
experimental approach emphasizing hypothesis testing, statis-
tical analysis, and development of models to provide new
insights and develop new hypotheses. A hypothesis is any state- '
ment that can be tested.
9. Two approaches to testing hypotheses are the inductive method,
which goes from the specific to the general, and the deductive
method, which goes from the general to the specific. The indue- .
tive approach is useful for investigating correlations between
classes of facts. In the deductive method, the investigator devel-
ops a research hypothesis, collects data to support or refute it,
develops a'mathematical model, attempts to fit the model to the
data, and then tests and, if necessary, modifies the model.
10. Testing hypotheses entails the collection of data by direct
observation or by experimentation. Experimentation involves
simplification by manipulating one or a few variables while
holding others constant. Both laboratory and field experiments
involve manipulations of one or afew independent variables in.
.11. Experimentation involves 'the determination of the response of
one variable, the dependent variable, to variations in an inde-
pendent variable or variables, mru:iii;nlated by treatments. The
association of the dependent to independent variables assesses
the nature of the response of the variable and tells
something about the relationship between the two. The exper-
. imental approach involves replicates of treatments and controls
that do not receive the treatment. Replicates allow the scientist
to account for uncontrolled variations among experimental
units; controls form the basis __esults of
experimentation allow the investigator to reject or accept the
hypotheses developed.
12. A model is an abstraction and simplification of a natural phe-
nomenon developed to predict a new phenomenon or to pro-
vide insights into existing ones. Averbal or graphic model may
serve as the basis of a more formal mathematical model. A
mathematical model may be statistical or nonstatistical. Non-
statistical models may be either analytical or simulation. Ana-
lytical models are mathematical formulations that can be
solved directly. Simulation models may take on a variety of
forms, including differential Because they cannot be
solved analytically, simulation models require the use of a com-
i puter to arrive at a solution.
13. Once developed, the model should be validated. Validation is
the test of the model's ability to do what it is supposed to do.
It measures quantitatively the extent to which the output of the
model agrees with the behavior of the real-life system.
Reyiew \ouestions
1. Define ecblogy.
2. Why was plant geography the apparent stimulus for the devel-
opment of modem ecology?
3. What differences separate the organismal concept of ecology
from the individmlli.stic concept?
4. How do the two concepts in question 3 relate to holism and
reductionism in ecology?
5. What is applied ecology and how does it relate to theoretical
and ecosystem ecology?
6. The following statement appeared in Sustainable Long-Term
Forest Health and Productivity by the Society of American
Foresters: " ... is the strategy by which in the aggregate, the full
array of forest values and functions is maintained at the land-
scape level. Coordinated management at the landscape level,
including cross ownership, is essential component." This state-
ment caused serious divisiveness among foresters. Why?
7. Refer to the document, ''The Sustainable Biosphere Initiative"
(Lubchenco et al. 1991). Select one of the research topics and
discuss how the results from such research would relate to our
environmental problems.
8. Why is ecology not taken as seriously as it should be by polit-
ical decision makers?
9. What makes ecology an empirical science?
10. What is a hypothesis?
11. How do the inductive and deductive approaches to testing
hypotheses differ? What are advantages and weaknesses of each?
12. What distinguishes a dependent variable from an independent
variable?
13. What is the importance of replicates and controls in ecological
studies?
14. What is a model? Why are models useful in ecological research?
15. What is the difference between statistical and nonstatistical
models?
16. Comment on the statement made by C. J. Krebs: "Hypotheses
without data are not very useful, and data without hypotheses
are wasted." What does Krebs mean?
17. Examine some papers in such ecological journals as Ecology,
Journal of Ecology, and Journal of Animal Ecology, both in
early and recent years. HQw do they differ in experimental
approach? Are the hypothese-s clearly stated'? Are the data col-
lected adequate for testing the hypotll:eses?