A Brief History of Ecology1 Humans as image bearers of their Creator, have been endowed with both the motivation

and the capacity to sense wonder, to experience curiosity, and to desire an understanding of the relationships with their environment. These human abilities were activated and used when Adam was given the first recorded assignment by his Creator. Each kind of creature was presented to the man to see what he would call them; and whatever the man called a living creature, that was its name. The man gave names to all the cattle, and to the birds of the sky, and to every beast of the field, but for Adam there was not found a helper suitable for him (Gen. 2;19-20). Given that naming in Jehovahs perspective always implies significance or relationship, the implication is that Adam studied the nature of each creature and its possible relationship to himself and the other creatures; and that he reached a point of understanding of how each related to him and to other creatures. With this knowledge, Adam was able to conclude that no other creature was able to relate to him as a human being. This new awareness prepared Adam to receive the woman God would create for him, and would also prepare him to exercise proper dominion and stewardship of the garden (Gen. 1:26-28; 2:15). We learn from these verses that God has equipped humankind to learn of His design and purpose for each living and nonliving part of His creation. Genesis 1:24 records, Then God said, Let the earth bring forth living creatures after their kind: cattle and creeping things and beasts of the earth after their kind; and it was so. Each kind of plant and animal was created with the purpose of multiplying after its kind and occupying particular habitats within the creation. This principle is emphasized in Psalm 104: 16-18 where we read (emphasis added):
The trees of the LORD drink their fill, The cedars of Lebanon which He planted, Where the birds build their nests, And the stork, whose home is the fir trees. The high mountains are for the wild goats; The cliffs are a refuge for the rock badgers.

Note that each creature is given a habitat within which to fulfill its purposes as part of the Creators plan for each of them. Each creature is seen as good and therefore has value which is defined apart from their value to mankind i.e. intrinsic value. Thus, each kind of creature is genetically endowed with the capacity to reproduce and to fit into a particular niche, including suitable habitat, food, shelter, and other resources necessary for it to prosper and experience Gods blessing (Gen. 1:22,28). Throughout history humans have demonstrated innate curiosity, desire to learn, and the rational capacity to understand our relationship with other creatures and with our physical surroundings. The study of relationships between organisms and their environment has come to be known as ecology from the Greek oikos (house) + logos (study of). However, the historical paths leading to its current status as a biological science are difficult to retrace. This summary is an attempt to summarize some of the notable developments and contributors.

John E. Silvius, Senior Professor of Biology, Cedarville University, Cedarville, OH. Compiled in 2007 from the listed references.

A Brief History of Ecology

Beginning around 320 BC, Theophrastus, a brilliant student of Aristotle and Plato began a series of studies that would earn him the name of father of botany. He classified over 500 plants into major growth forms trees, shrubs, herbs; and further organized them according to morphology. Theophrastus is also regarded by some as the founder of ecology. He conducted seed germination experiments and discussed the influence of abiotic habitat factors on plants. In particular, he studied the spatial and temporal variations in the vegetation of flood plains along rivers. The next wave of progress in the history of ecology was spurred by the Renaissance scientists of the 16th and 17th centuries. Many contributions of Aristotle and other Greek philosophers continued to be influential well into the 18th century. Thus, Aristotles scala naturae or Great Chain of Being became the unifying framework of nature in which each element was assigned a place level within the ladder of life. Artists chose plants, animals, and landscapes as their subjects. Natural history emphasized the study of creatures in relation to their natural environments using a descriptive approach; whereas, natural philosophy, the forerunner of physics and chemistry, emphasized a quantitative and conceptual approach to understand the natural world. Natural theology dealt with the questions of origin and purpose of the natural world. Its proponents believed that the natural world had the marks of design and purpose (the teleological argument). Thus, they were at odds with the mechanistic philosophy of the day (i.e. view of nature and organisms with a machine metaphor). Swedish naturalist, Carl Linnaeus, (17071778) became a prominent influence in the 18th century. His expertise in natural history and natural theology led him to the notion that biotic interrelationships were designed by God to work harmoniously and permanently and for the benefit of humanity (Edgerton, 2007). Although early in his career, he believed that God had created each species, he later revised his concept to suggest that God had created only a relatively few kinds of organisms from which hybridization then gave rise to many more species. Linnaeus and his students extensively researched such topics as the food preferences of grazing animals (2,314 experiments), the number of grazing animals sustainable by a given pasture, the survival and dispersal of seeds through animal dung, and the pathology of parasitic worms. His Politia Naturae (1760), On the Police of Nature, described his theory of how populations are kept in check so that many species can share a given environment. Linnaeus was addressing the concept of ecological diversity: each species is confined to its own station (habitat) (Edgerton, 2007). He estimated that Sweden had about 1300 plant species, but only about 50100 are in any one place. Linnaeus argued that although that plants were created for the use of animals, actually, animals were created to regulate plants abundance. As proof, he cited numerous insect species that only eat a single plant species, doves eat surplus seeds, and other birds, bats, and anteaters eat insects to prevent them from consuming all of the plants they eat, and so on.

A Brief History of Ecology

Linnaeus used the term oeconomia naturae (1749), or natural economy, to mean what we now would call physiology, the study of how organs contribute to the functioning of the whole. He stated: By the Oeconomy of Nature we understand the all-wise disposition of the Creator in relation to natural things, by which they are fitted to produce general ends, and reciprocal use. [Note the emphasis on teleology and design, the themes of natural theology in Linnaeuss day; and which would soon be articulated by William Paleys watchmaker analogy in his book, Natural Theology (1802), and often cited in intelligent design arguments today.] Linnaeus oeconomy of nature, perhaps partly derived from his knowledge of Scripture (Gen. 3:19 For you are dust, And to dust you shall return, and Isa. 40:6 All flesh is grass.) included a theory of the cycling of nutrients before the chemistry of mineral elements and gases was discovered (nitrogen by Rutherford, 1772; oxygen by Priestley, 1771). Linnaeus explained:
when animals die they are converted into mould, the mould into plants. The plants are eaten by animals, thus forming the animals limbs, so that the earth, transmuted into seed, then enters mans body as seed and is changed there by mans nature into flesh, bones, nerves, etc.; and when after death the body decomposes, the natural forces decay and man again becomes that earth from which he was taken.

While Linnaeus influence was spreading, a Prussian botanist, Carl Ludwig Wildenow, was studying changes in plant species over time in peat bogs. This dynamic series of changes in the environment and in plant and animal populations over time has come to be known as ecological succession. Like Linnaeus, Wildenow traveled extensively. A major contribution was his proposal that similar climates supported similar vegetation. One young naturalist who was intrigued by Wildenows ideas was a young and wealthy Prussian student named Alexander von Humboldt. At the age of 30, in 1799, Humboldt and a French botanist, Aime Bonpland embarked on a 5-year expedition across Central and South America and the Carribean islands. His extensive study, plant collections, and cataloging of field observations led Humboldt to propose that plant species are distributed in associations that are predictable when one characterizes the environmental conditions of the geographic location. Using astute observations and inductive logic (moving from specific observations to generalization) Humboldt was able to combine the descriptive approach of natural history and the quantitative and conceptual approach of natural philosophy to lay the foundation for biogeography. Humboldts contribution to the development of ecology and other sciences was further developed through his speaking and writing. His five-volume work, the Kosmos, was his successful attempt to unify various branches of science and philosophy into a comprehensive description of the physical world. Humboldt is regarded by many as a giant of the 19th century and perhaps the last individual that came close to presenting a comprehensive treatise of the knowledge of the world in his time.

A Brief History of Ecology

Following Humboldts lead, a stream of naturalists with the interest in biogeography traveled to the western hemisphere. Danish botanist, Johannes Warming (1841-1924) spent three years in Brazil and expanded upon the studies of Humboldt by describing the effects of temperature, moisture, soils on patterns of vegetation. His influence upon biogeography spread through his textbook Plantesamfund, which was later translated to English with the title, The Oecology of Plants: an Introduction to the Study of Plant Communities. In it, Warming introduced his classification of plants based upon their adaptations to moisture and salinity; namely, hydrophytes, mesophytes, xerophytes, and halophytes. Biogeographical studies such as Warmings stimulated curiosity about how plant growth is specifically affected by environmental factors. Justis von Leibig (1803-1873), a German chemist and colleague of Humboldt, is regarded as the father of the fertilizer industry for his discovery of nitrogen as a required plant nutrient. His research demonstrated that incremental increases in nitrogen would cause increases in plant growth. From these data, he proposed what has come to be known as the law of the minimum; namely, that the rate of a process is controlled by the factor in shortest supply. Later, F.F. Blackman (1905) expanded the Leibigs law to include a corollary that a process is inhibited by excessive levels of an otherwise necessary factor. These concepts became the foundation for the discipline of physiological ecology. Animal ecology developed much later than plant ecology and from different roots. Animal ecology was influenced by another naturalist, Charles Darwin (1809-1882), who himself was influenced by Humboldt. Like Humboldt, Darwins thinking was shaped by his travels to South America and nearby islands, the Galapagos. He collected specimens extensively and wrote voluminous field notes, comparing animal life on the continent with that of the islands. From these observations and from the influence of Thomas Malthus, an economist, Darwin proposed his theory of population growth, inevitable resource limitation, and survival of the fittest as a mechanism of natural selection and biological evolution. From Darwins work and the contribution of his unknown contemporary, Gregor Mendel (1822-1884), whose quantitative theory of inheritance became the foundation of genetics. When Mendels work was discovered shortly after 1900, natural selection came to be defined quantitatively as changes in gene frequency within a gene pool shared by all interbreeding members of the population. Out of these concepts grew the science of population genetics. Other scientists combined the Malthusian concept of population growth and limitations in studies of population dynamics in specific environments. Out of these studies and the work of two mathematicians/physicists, Alfred J. Lotka (USA) and Vito Volterra (Italy), in the 1920's came population ecology, a quantitative approach to ecology that emphasized the effects of interspecific competition and predation. In the late 1800's, as ecology was beginning to emerge as a biological science in its own right, a rift occurred between the plant ecologists and the animal ecologists. Plant ecologists voted to drop the o from oecology at a Botanical Congress, in Madison, WI in 1893. Zoologists would have nothing to do with the new term, fearing that botanists would usurp the discipline entirely. The fear was partly justified because some plant ecologists ignored the

A Brief History of Ecology

interactions between plants and the animal parasites and herbivores. So, animal ecologists considered adopting the term ethology which had already grown from zoological studies of animal behavior e.g. Conrad Lorenzs studies of animal imprinting. The emphasis of ethology was upon the explanation of differences in character from psychological theory (ethos = custom, or character). Before the schism completely separated the botanists and zoologists, two individuals published major works that emphasized the interrelationships between plants, animals, and environment. The American plant ecologist, Frederick E. Clements (1874-1945), influenced by Humboldts plant association concept, proposed the concept of a biotic community in which plants and animals have equally important roles. Clements colleague, Victor E. Shelford (1877-1968), a zoologist at University of Illinois had been studying the response of animal populations to environmental changes. Shelford drew from the work of Leibig and Blackman to conceive his law of tolerance; namely, that individuals and populations have a tolerance range between minimum and maximum supply of each environmental factor and within which they will function optimally. Together, Clements and Shelford, authored a textbook Bio-ecology, in which they introduced the concept of biomes representing geographic areas containing a distinct flora and fauna as influenced by particular climate conditions. This dynamic duo served not only to unify botanists and zoologists under an ecological umbrella but also forged a strong union between biogeography and physiological ecology, the latter providing a physiological basis for plant and animal distribution. However, Clements the unifier soon became the source of another schism in the fledgling ecology. Clements studied ecological succession in plant communities and began to view the community as a dynamic system that functions like an organism (or superorganism) which moves predictably through changes from a youthful stage colonizing bare ground to a mature, self-sustaining climax in which the community is at equilibrium with the physical environment. If disturbed, the community can retrace its ontogeny from youth to maturity again, just like an individual organism. Clements view of community-as-organism was based upon a holistic philosophy in which the whole is viewed as greater than the sum of the partsthat the whole contains emergent properties, new properties of the system (e.g. community) that would be missed even if one fully understood the workings of each part (e.g. population). In 1926, Henry A. Gleason (1882-1975) challenged Clements organismic concept with the publication of The Individualistic Concept of the Plant Association. In the article, Gleason maintained that communities do not self-reproduce and develop as if they were organisms. Instead, each plant community is unique and arises from random processes (e.g. seed dispersal, fire, physical environment). In place of sharp boundaries delineating a community, each population exists along an environmental continuum and responds individually in such a way that

A Brief History of Ecology

determines its spatial distribution. Gleasons individualistic view of the community represents a reductionist philosophy that rejects the concept of emergent properties. Communities are composed of individual parts (i.e. populations), each of which interact with environmental conditions and other populations without the influence of an organizing principlethat moves the whole community through some developmental path. Communities therefore can be understood by studying the parts, because the whole is equal to the sum of the parts. Arthur G. Tansley (1871-1955) was an English botanist that for awhile was drawn to Clements organismic view but eventually rejected it. Instead, he coined the term ecosystem which he defined as all of the populations in a given environment (as components) and the physical environment included. Thus, Tansley retained a holistic approach but moved the whole to a higher hierarchical level, emphasizing that ecosystems are functional units that must be studied as systems. Ecosystem ecology has developed by ecologists such R. A. Lindeman (1942) who studied the trophic relationships as affected by long-term changes in Cedar Bog Lake, Minnesota. The Odum brothers Eugene, (1913-2002) and Howard (19242002) applied general systems theory to ecology with computer analysis of large amounts of data, giving rise to systems ecology. Even before Tansley, several biologists were using a holistic ecosystem-type approach. August Friedrich Thienemann (1882-1960), a German aquatic ecologist, published 460 papers on his research in aquatic communities with support for "super-organismic unity" but with an ecosystem approach through his emphasis on interactions between aquatic populations and the environment. Thienemann introduced the concept of nutrient cycling among populations which he classified as producers, consumers, and decomposers. Edgar N. Transeau (1875-1960) sought to improve crop yield efficiency by improving the energy trapping process of photosynthesis. He made estimates of energy transfer in experimental plots of corn on the campus of University of Illinois. Transeaus classic paper, The Accumulation of Energy in Plants (1926) introduced the concepts of energy budgets and net primary production by ecosystems. Transeau was chair of the Department of Botany at The Ohio State University until 1946 and turned it into a first-rate program. He proposed the Prairie Penninsula concept which invokes a post-glacial xerothermic period that caused the migration of prairie communities into Ohio. By the dawn of the 20th century, it became evident that the growing conversion of land and resources to human usage was threatening the soil, water, plant, and animal populations. Both ecologists and popular writers became influential to public opinion. Ecologist Paul Sears (18911990), wrote Deserts on the March in response to the effects of overgrazing and plowing which culminated in the 1930's dust bowl.

A Brief History of Ecology

Aldo Leopold (1887-1948) became a one-man department of wildlife management at University of Wisconsin, in 1924. His A Sand County Almanac has become a classic account of the Leopold family experience in restoration of a degraded farm and the birth of his land ethic. In 1962, Rachel Carson (19071964), a zoologist and marine biologist, published Silent Spring, which contained a sobering expression of her concern that the growing use of chemical pesticides was destroying bird populations. Her influence generated legislation restricting the use of the insecticide, DDT, and heightened awareness of toxic substances in the environment. The influence of articulate persons such as Also Leopold and Rachel Carson gave rise to the modern environmental movement. Ecology has become a household word since the 1960's because of this movement. Meanwhile, the science of ecology continues to maintain a scientific emphasis through professional meetings and publications of societies such as the Ecological Society of America, and through The Nature Conservancy, a scientifically sound organization of professional ecologists and laymen who work for the conservation of natural areas and biodiversity around the globe. Additional fields have spun off from the ecological sciences. Restoration ecology attempts to apply principles of ecosystem ecology to management and restoration of degraded land and aquatic systems in an effort to restore them as nearly as possible to their pre-degraded state. Conservation biology attempts to apply both ecological principles and an ethical foundation in effort to conserve plant and animal species. However, today ecologists are becoming more aware that it will take more than scientific data to convince society that human abuse of the Earth is pushing the planet toward destruction. Perhaps we have come too far from the Garden of Eden in which our father, Adam was given instruction and responsibility to serve God and steward His creation. Some Christian ecologists such as Calvin B. DeWitt of University of Wisconsin have emerged as proponents of an environmental stewardship ethic based upon Genesis 2:15 and other passages. DeWitt and others have established constructive dialog with secular biologists and ethicists who are recognizing that science needs the support of religious leaders. For example, secular environmental ethicist, J. Baird Callicott wrote:
The Judeo-Christian Stewardship Environmental Ethic is especially elegant and powerful. It also exquisitely matches the requirements of conservation biology. The Judeo-Christian Stewardship Environmental Ethic confers objective value on nature in the clearest and most unambiguous of ways: by divine decree.

A Brief History of Ecology

Most recently, Edward O. Wilson (1929- ), a distinguished biologist from Harvard University, co-author with Robert MacArthur (1930-1972) of the noted theory of island biogeography, and proponent of sociobiology and scientific humanism (he calls it "the only worldview compatible with science's growing knowledge of the real world and the laws of nature), has written an appeal to the evangelical church to to bring the theological and moral arguments for saving the Creation into partnership with science in an effort to avoid further environmental deterioration on planet Earth. Indeed, there would appear to be an openness for dialog between science and religion that is unprecedented. Such is a brief summary of the history of ecology. And, such are the exciting times in which we begin our study of ecology. May your encounter with this brief history and your acquaintance with a variety of personalities and philosophies stimulate you to engage in deliberate effort to make your study a fruitful step toward development of your profession in biology and in your vocation.
References:
Clements, F.E. and V.E. Shelford. 1939. Bio-ecology. McGraw-Hill. New York. Carson, R. 1962/2002. Silent Spring. Houghton M ifflin, Mariner Books. DeW itt, C.B. 1998. Caring for Creation: Responsible Stewardship of God's Handiwork. Baker, Grand Rapids, MI. Gleason, Henry A. 1926. The Individualistic Concept of the Plant Association. Bulletin of the Torrey Botanical Club 53: 7-26. Smith, R.L. and T.M. Smith. 2001. Ecology and Field Biology, 6 th ed. Benjamin Cummings. San Francisco. Egerton, F.N. 2001-2007. History of the Ecological Sciences. Bulletin of the Ecological Society of America. http://www.esapubs.org/bulletin/current/history_links_list.htm History of Biology. 2007. Wikipedia. http://en.wikipedia.org/wiki/History_of_biology#Natural_history_and_natural_philosophy Leopold, A. 1948. A Sand County Almanac. Oxford Press. Lotka, A.J. 1925. Elements of Physical Biology. Williams and W ilkens, Baltimore. [Reprinted by Dover Publ. 1956. as as Elements of Mathematical Biology.] W ilson, E.O. and R.H. MacArthur. 1967/2001. The Theory of Island Biogeography. Princeton University Press. W ilson, E.O. 2006. Creation: An Appeal to Save Life on Earth. W .W . Norton. W orster, Donald. 1994. Nature's Economy: A History of Ecological Ideas. Cambridge University: Cambridge.