Community Ecology History of community ecology Outline: 1. History of community ecology from pre 1700s to today A. pre-1700s: an unordered and mystical world B. 18th century: descriptive (natural history) studies C. 19th century: "ecology" coined, Darwinism D. late 19th/early 20th centuries: self-awareness of ecology, "community" coined E. 20th century: Clements/Gleason succession debate, contributions from MacArthur 2. Early emphasis was on pattern; more recent research has explored process Terms/people: Haeckel von Humboldt Darwin Forbes Möbius Cowles Lack superorganism uniformitarianism (Lyell) Clements Gleason Elton individualistic MacArthur Hutchinson Shelford oikos Ecology has proceeded rather like a relay race, with each “leg” of the race differing from others in terms of pace, trajectory, and influence. The first major leg of ecology came from the age of exploration, with repeated patterns of species co-occurrences being noted. The second leg of ecology’s history is from late 19th-century/early 20th-century physiologists who tried to determine form as a consequence of environment. The third leg came from an emphasis on biotic interactions (especially competition). The fourth leg witnessed a greater emphasis on discerning process and not merely documenting pattern, comparing patterns with null models, and being skeptical of competition as the main driving force behind communities. Some may argue that we are now in the fifth leg, focusing on the importance of taking scale into account, applying community ecology to conservation, and branching out into new directions. Roots of CE: natural history, plant geography -"balance of nature" -3 pathways of development and focus: 1. Response physiology - physiological limitation of plants and animals by environmental factors - Haeckel, later Cowles 2. An evolutionary theme - Darwin 3. Plant geography - focus on mapping associations and vegetation formations (what we would now call communities) - von Humboldt pre 18th century: -biology largely unordered (pre-Linnaeus) -world still highly mystical 18th century: -natural history - descriptive -order/progress (influenced by the Church) -equilibrium -Linnaeus - orderly scientific nomenclature (hierarchical) -Buffon - naturalist, first described how environment shapes animals So, prior to 19th century: emphasis was on describing patterns. 19th century: early developments were in Europe -Alexander von Humboldt 1807 -early 1830's: Charles Lyell - principle of uniformitarianism -1838: P.F. Verhulst developed formula for population growth when resources are limiting -1859: Charles Darwin’s Origin of Species: scientific revolution from natural selection -Ernst Haeckel 1866 late 19th/early 20th centuries: -crystallization of ecology, self-awareness of ecology -Karl Möbius 1877 (translated in 1883) -Stephen A. Forbes 1887 -Johannes Eugenius Bulow Warming 1896 - "Oecology of plants: an introduction to the study of plant communities" - first to ask coherent questions that still dominate community studies: 1. Why does each species have its own habitat? 2. How do species congregate to form characteristic communities? - recognized importance of dominants and subdominants, adaptive value of lifeforms, effects of fire on community composition and succession, importance of phenology concluded that soil has more of an effect on vegetation than climate, and emphasized temperature and moisture as prime climatic factors. -Henry Chandler Cowles 1899 - U. Chicago, succession, stability; studies changes in plant community on the sand dunes of Lake Michigan between 1898 and 1911. Emphasis on the dynamic nature of vegetation. 20th century: -Victor Shelford 1915 - U. Nebraska, first president of ESA (founded 1915); looked at animal communities in the same manner that plant ecologists did -Frederic Clements 1916, 1936 - U. Nebraska, one of the most influential ecologists "superorganism" -Henry Gleason 1917, 1926 - IL Nat. Survey Individualistic view of community structure -Clements vs. Gleason debate of whether a community is a definable unit vs. a level of organization above that of populations in a given area -Charles Elton 1927 - animal ecologist, niche, food webs; book Animal Ecology (1927) written in a mere 85 days and is still in print -David Lack 1940s – role of interspecific competition in shaping community structure -1940-78 - G. Evelyn Hutchinson - limnologist and zoologist; niche; Ph.D. advisor for Robert MacArthur -1950s-1960s - Odum brothers (Howard Tom and Eugene P.) - begin ecosystem studies, synthesize relatively new ideas on ecosystem ecology -Robert H. MacArthur 1950s-60s - Hutchinson’s student; birds, very thoughtful and creative ecologist who tragically died young (in 1972 at age 42) but who formulated/influenced current thinking on competition, niche and habitat partitioning, biodiversity, food webs, extinction, importance of evolutionary life history traits, conservation biology and reserve design, and other topics; most influential ecologist in modern history -influence of non-English-speakers (e.g. Vasily Dokuchaev) -1968-1974 - International Biological Program (IBP) - $27 million spent on large-scale ecology research projects; up to this time, very little money was spent on ecological research -1980 - National Science Foundation (US) establishes Long-Term Ecological Research sites (LTER) Current approaches in community ecology: 1. Community classification, derived directly from the Clementsian view; in Europe this led to rigid typological classification schemes, whereas in North America the emphasis was more on quantitative approaches with more of a Gleasonian perspective. 2. The holistic community view of Clements mutated into the ecosystem/energy ideas of Eugene Odum. But ecosystem ecology had its formal beginnings in the mid-1930's when Arthur Tansley coined the term (in response to the excesses of a rigid Clementsian view). This, in turn, was picked up by Raymond Lindeman, who developed the "trophic-dynamic aspect of ecology" (i.e., formalized the routes of nutrient and energy transfer). Ecosystem ecology later developed along two pathways: systems ecology and biogeochemistry. Even today, ecosystem and community ecology may be taught together as synecology, reflecting their shared history. 3. Meanwhile, the evolutionary foundation of Darwin influenced Charles Elton’s thinking about ecological niches and adaptations of organisms to their environments. Elton and Gause (who developed mathematical approaches to multi-species dynamics) in turn influenced G. Evelyn Hutchinson and David Lack. Together, they affected the thinking of Robert MacArthur, who championed a mathematical/theoretical approach to communities in the 1950’s and 60’s. MacArthur’s influence is still pervasive through much of what community ecologists do today. 4. The individualistic approach of Gleason was the foundation of the thinking of the Australian ecologists Harry Andrewartha and Charles Birch, whose 1954 book, The Distribution and Abundance of Animals, established a framework for an individual-based approach to communities. There have been various attempts to define the most important concepts or central themes of ecology. Here are what I believe to be the three central themes of contemporary community ecology: 1. Ecological systems are variable in space and time (cf. the equilibrium/homogeneity paradigm). 2. Ecological patterns and processes are scale-dependent (cf. scale is irrelevant or too messy). 3. Ecological systems are open to external influences (cf. the closed-system paradigm). Next time: ecology as a science, the scientific method, hypothesis-testing References: Gleason, H.A. 1926. The individualistic concept of the plant association. Bull. Torrey Bot. Club 53:7-26. Golley, F.B. 1993. A History of the Ecosystem Concept in Ecology. Yale University Press, New Haven, CT. Hagen, J.B. 1989. Research perspectives and the anomalous status of modern ecology. Biology and Philosophy 4:433-455. Hagen, J.B. 1992. An Entangled Bank: The Origins of Ecosystem Ecology. Rutgers University Press, New Brunswick, NJ. McIntosh, R.P. 1985. The Background of Ecology. Cambridge University Press, Cambridge, UK. McIntosh, R.P. 1995. H.A. Gleason’s ‘individualistic concept’ and theory of animal communities: a continuing controversy. Biological Reviews 70:317-357. Odum, E.P. 1977. The emergence of ecology as a new integrative discipline. Science 195:12891293. Sheail, J. 1987. Seventy-five Years in Ecology: The British Ecological Society. Blackwell Scientific Publications, Oxford, UK.