2 Lesson Brief history of plant ecology
Key historical personalities Major phases in the development of plant ecology concepts
European biogeographical phase European plant community classification phase American concepts of succession, continuum, and individualistic response of plants Ecophysiology, population dynamics, and ecosystem modelling phase Big ecology and current focus on global change
European biogeographic phase
Plant ecology roots in the observations of the great variety of form in nature and the desire to develop a consistent means of describing the variety of vegetation forms on the Earth. Observation that similar climates produce similar plant forms.
Betula pubescens forest, Finland
For example, cactus growth forms common to many hot deserts.
European plant geographers
Explored the worlds biomes seeking to develop a consistent means to describe the diverse plant life of the globe. Von Humboldt traveled to Central America,
Student of Carl Ludwig Wildenow. Measured environmental parameters Collected 60,000 plants Wrote a 30 vol work (Vayage aux Regions Equinoxiales), Correlated environmental factors with vegetation. Noted that similar climates produced similar vegetation, even in very disjunct regions. Biography: Botting, Douglas. 1973.
Humboldt and the Cosmos. New York: Harper and Row.
Alexander von Humboldt (1773-1858)
Aim Bonpland (1773-1858)
Von Humboldts botanist in S. Amer. Attempted to complete the 17 vol. botanical studies from the expedition, but only completed 4 vol. Remaining volumes were completed by Gottleid Christian Kunth. Settled in S. America, was jailed by Paraguan dictator for 10 years and finally moved to Uraguay and gardened until he died.
Humboldts Orinoco R. Expedition (1799-1804)
Orinoco R. and the Casiquiare Canal to Esmeralda
Expedition camp on the Orinoco R.
Cinchona condaminea (Quinine) From Plantes equinoxiales
Chimboraza, Equador (20,702_
Humboldt and Bonpland nearly reached the summit (19,292 ft.), highest point reached by westerners to that time and the record stood for 30 years. Climb was the most famous accomplishment of the expedition. Vegetation zonation diagram was also a major accomplishment, Presaged later elevation gradient diagrams by Merriam and Whittaker.
Botanical Heritage of the South American Humboldt Expedition
60,000 specimens 3,000 new species to science 17 lavishly illustrated volumes
Inga excelsa from a volume devoted to Mimosas and other leguminous plants.
Other famous European early plant geographers
J.F. Schouw (1789-1852)
described the role of environmental factors on plant distribution in 1823, emphasizing the role of temperature, popularized the idea of using suffix -etum for naming associations.
Transplant gardens in Tyrolean Alps (300 species). Distinguished inheritable from environmentally-affected factors. Beautiful vegetation descriptions and understanding of succession in :
Anton Kerner von Marilaun (1831-1898) (Kerner)
___________. 1863. Plant life of the Danube Basin. Translated by H.S. Conrad. 1951.
Succeeded Willdenow at Gttingberg. Described more than 50 major vegetation formations worldwide in modern physiognomic terms, relating their distribution to climatic factors.
August Grisebach (1814-1879)
Johannes Eugenius Blow Warming (1841-1924) Andreas Franz Wilhelm Schimper (1856-1911)
Johannes Eugenius Blow Warming (18411924)
First true ecologist Studied a tropical woodlandsavanna in Brazil for 3 years. 30 years later wrote: The Oecology of Plants (1892 in German and an English edition in 1906). London: Oxford University Press; which is a model for modern ecological studies today; Organized the world's first ecology course and lecture notes were published as the first plant ecology text; Synthesized plant morphology, physiology, taxonomy, and biogeography into a coherent science for the first time; Among his findings: (1) soil has larger impact on vegetation than climate, (2) emphasized moisture and temperature as prime climatic factors; (3) coined many ecological terms still used today (e.g. halo-, hydro-, meso-, xerophyte).
Andreas Schimper (1856-1901)
University of Bonn. Plant Geography on a Physiological Basis (1903). He stressed morphological features of presumed adaptive value. Dwight Billings 50 years later called him the father of plant physiological ecology. Stressed climatic and edaphic factors. Borrowed heavily from Warming, but did not acknowledge him.
Illustrations from Pflanzen-Geographie
Christian Raunkiaer
University of Copenhagen. System of botanical lifeforms, a categorization based on the position of the wintering buds. Raunkir (1934), The Life Forms of Plants and Statistical Plant Geography.
Raunkiaers growth forms
European plant community classification phase
In Europe in the early 20th Century, most vegetation scientists were occupied by classification of vegetation. Most famous was Josias BruanBlanquet (1884-1980).
Followed in path of Kerner. Developed methods of community sampling, data reduction, association nomenclature. Cooperated with Rbel in Switzerlandl to develop an approach to plant synecology (plant community classification) called the Zurich Montpellier School of Phytosociology. He founded a research station at Montpellier, France called the Station Internationale de Geobotanique Mediterrenne et Alpine (SIGMA).
Josias Braun-Blanquet (1884-1980)
Braun-Blanquet sorted-table analysis method
This is a sorted table for the shoreline plant communities in the Netherlands.
Table analysis methods
A fundamental aspect of the vegetation analysis performed in Montpellier School of phytosociology is the use of sorted tables, whereby diagnostic plant species are identified for plant associations. Other similar schools of vegetation classfication developed in northern Europe under Du Rietz. In England vegetation ecology developed under the guidance of Sir Arthur Tansley who founded the British Ecological Society, coined the term ecosystem, and called for more physiological investigations in field studies.
Heinz Ellenberg (1913-1997)
Taught with Heinrich Walter at the University of Stuttgart. Demonstrated the differences between the physiological and ecological behavior of plants. Vegetation Ecology of Central Europe: one of the highest expressions of the European approach to vegetation description. Also wrote the classic textbook: Aims and Methods of Vegetation Ecology with Dieter Mueller-Dombois
Russia and U.S. ecology: Huge landscapes
A colluvial basin in northern Alaska
Contrast between Europe and North America in the early 1900s
In the US, and in Russia in the late 1800s and early 1900s ecologists became fascinated the large landscapes of the north and noted the repeating assemblages of plants on large landscapes. Vegetation ecologists became interested in the spatial patterns of vegetation associated with larger landscapes and the response of plants along long environmental gradients, in contrast to the transitions seen along abrupt boundaries such as those described in the Alps by BraunBlanquet and the European phytosociologists. This previous shows an area in the northern foothills of Brooks Range, Alaska. It is an example of the dominance of natural landscape patterns, in contrast to the small-scale patterns created by humans that dominate many European landscapes and now most of the USA in the lower 48 states.
Two key Eastern European ecologists whose ideas predated those of famous ecologists in North America:
Jozef Paczoski (1864-1941)
Considered by many to be the Father of phytosociology; showed how plants modify the habitat, create own microenvironment; discussed role of competition; causes of succession, role of fire, interdependence of species in communities; graphs of direct ordination (1930) were the first such analysis; founded world's first department of phytosociology in Poznom, Poland.
Leonid Ramensky (1884-1953)
Preceded ideas of Gleason and Whittaker (individuality of species and continuum of vegetation); coined the term phytocoenosis; expressed community composition in a table form much like Braun-Blanquet; prefigured r-K and C-S-R categories of Grime, MacArthur, and Pianka, He noted three categories of plants: violent (= competitors, K-strategists); patient (stress-tolerators) and exploring (ruderals, r-strategists).
Clinton Hart Merriam (1955-1942)
M.D. from Columbia U. Chief of the U.S. Biological Survey (1885-1910). Had a large influence on the early development of plant ecology and concepts of vegetation in relation to climate. Led large expeditions through the west and developed ideas of elevation and latitudinal "life-zones", which were later disputed mainly by Daubenmire.
Merrriams life zones
Based largely on a biological survey of San Francisco Peak area (Merriam 1890). Latitudinal and altitudinal zones were based primarily on summer mean temperature and secondarily on humidity, but zonations had little empirical data to support them. His concepts of elevation zonation show some similarity to those of Humboldt in the early 1800s.
Henry Chandler Cowles (1869-1939)
QuickTime and a TIFF (Uncompressed) decompressor are needed to see this picture.
QuickTime and a TIFF (Uncompressed) decompressor are needed to see this picture.
Geologist-turned-botanist. Heavily influenced by the ideas of Warming. Was struck by the relationships between the landforms, geology, and vegetation. One of the founders of the school of dynamic ecology with studies of succession. Most famous for his studies of succession on sand dunes around Lake Michigan (1899, The ecological relations of the vegetation of the sand dunes of Lake Michigan)
American concepts of plant succession
Pond succession in Ohio based on the work of Dachnowski. Based on ideas of Clements and Henry Cowles Classic diagrams, such as the one at to the left, were produced by the followers of Cowles. Many students at the University of Chicago including Gleason, William Cooper, Edgar Transeau, and Emma Lucy Braun. Helped organize the Ecological Society of America in 1911, and became president of ESA in 1918.
Frederick Edward Clements (1874-1945
Perhaps the most famous of the early American ecologists. Born, raised, and educated in Nebraska. Enchanted with the stories of Lewis and Clarks journey across North America in 1803-06 and made long journeys of his own throughout N. America. Focused on the role of climate in shaping the vegetation and the causes of succession. Wrote the Phytogeography of Nebraska (1899) and later described much of the vegetation of North America, naming regional formations, associations and seral stages.
Clements detailed observations of plants and plant communities
In 1917 he became associated with the Carnegie Institute's coastal laboratory at Santa Barbara and the alpine laboratory on Pikes Peak, and in 1929 he wrote his famous plant ecology textbook with John Weaver. Described the methods of vegetation sampling, and vegetation and strategies of plants in most of the biomes of the west. This diagram is from that book and shows the rooting strategy of buffalo grass.
Clements journeys
Clements traveled all over the west and saw the vegetation communities as "organisms" that followed prescribed life-cycles that all ended in the climatic climax. This organismic approach was disputed--rather quietly by many prominent ecologists, and rather heatedly by Henry Gleason. The early days of plant community ecology were ones of cataloging and describing plant communities with respect to patterns in the landscape, patterns that were obvious at a human scale.
Henry Gleason (1882-1975)
Student of Henry Cowles; primarily a taxonomist, with a strong grounding in the 'physiographic ecology' Cowles. of Wrote about the continuous nature of vegetation. Though of plant associations in terms of floristic composition. He noted that most species occurred along gradients of environmental gradients and that plant communities were composed of plant species, each of which was reacting individualistically to these gradients.
Gleasons individualistic hypothesis
Disputed Clement's organismic approach and developed the "individualistic hypothesis", whereby species responded individualistically to environmental conditions and the vegetative community did not act as an organism in any sense of the word. Studied the Mississippi Valley and the changes in community composition along the length of the river. Gleason was considered a heretic and a "good man gone wrong" from 1926 until the late 1940's and early 1950's when Cain, Curtis, Whittaker and others began developing ideas of the continuum approach.
John T. Curtis (1913-1961)
University of Wisconsin, professor of botany. Plant physiologist, orchid specialist. Strongly influenced by Gleasons individualistic hypothesis. Leading proponent of the continuum concept. Developed indirect ordination methods. He and his graduate students developed quantitative point sampling procedures and methods of ordination. Created the Wisconsin School of ecological thought, which strongly influenced the direction of American ecology up to the present. 39 Ph.D students, including Grant Cottam, Bob McIntosh, David Archibald,, John Bray, Orie Loucks, and Bob Burgess. He and his students published The vegetation of Wisconsin: An ordination of plant communities.
Mathematical approaches to analyze vegetation
Tilia americana
Curtis recognized that species were distributed individualistically along environmental gradients. He saw plant communities as composed of species coming and going along environmental gradients. He developed ordination techniques that portrayed vegetation communities relationship to each other based on their floristic similarity. Diagrams such as this one show species distributions in response to mathematical gradients that could then be correlated with environmental factors.
Bray and Curtis Ordination
Robert H. Whittaker (1920-1980)
Probably the most famous vegetation scientist in the US. His fields of interest included classification of communities, development of ordination and gradient analysis, measurement of species diversity and assessment of its significance, studies of driving forces of succession, comparative studies of biomass and plant productivity, analysis of roles of inhibitory metabolic compounds (allelochemicals). Produced 5-kingdom classification for organisms.
Gradient analysis of Smokey Mountain ecosystems
Like Curtis, Whittaker, developed methods to portray vegetation along complex environmental gradients. Primarily examined vegetation along elevation gradients in mountain ecosystems (Smokey Mountains, the Siskiyou Mountains of Oregon, and the Santa Cantalina Mountains in Arizona). Developed diagrams like this one which shows the distribution of a plant species, red maple, within the complex elevation gradients of the Smokey Mountains.
Ecophysiology Phase
About the same time that Curtis and Whittaker were having a major impact on plant community ecology, a group of ecophysiologists were also having a major impact in the 1940s and 1950s. Dwight Billings was one of the major proponents of using an ecophysiological approach in arctic and alpine systems.
Studied the autecology of a wide variety of alpine plants, and was a major proponent of using mesotopographic gradients, to study the response of plants along moisture gradients. Studied the effect of substrate on plant growth and success, ecological races, plant metabolic rates, and the effects of temperature and moisture stress on plants.
Dwight Billings (1910-1997)
Another key modern ecophysiologists in the Arctic is Terry Chapin here at UAF.
The newest plant ecology textbooks are primarily studies in plant physiology
Schulze, E.D., E. Beck, K. Mller-Hohenstein. 2002. Plant Ecology. Berlin. Springer.
First chapter is on stress physiology in plants.
Crawley, M.J. (ed.) 1997. Plant Ecology. Oxford: Blackwell.
First chapter is on photosynthesis:
The growth of plants depends upon their capacity to incorporate atmospheric carbon into organic compounds through the use of light energy absorbed during photosynthesis. This is a two-step process: (i) an initial photochemical reaction traps light energy in absorbing pigments (chlorophyll and accessory pigments).
Population biology
The study of plant populations is also a relatively new event. Although Clements did a few studies of plant competition and Tansley studied populations of Galium on different soils, the study of population biology was almost solely a topic of zoologists until John Harper really created the field in the late 1940s and 1950s after he met Charles Elton, a famous animal ecologist noted primarily for his studies of small mammal populations. Harper made the field of plant population biology quite accessible through this book which was first published in 1978.
Idealized plant history (Harper 1977)
This idealized diagram developed by John Harper shows the major stages in a plants life. 1. Starting with seed pool (the dormant phase), some of the seeds will not sprout and become seedlings due to various factors, such as unfavorable site, seed herbivory, or climate (the environmental sieve). 2. Of the seedlings that sprout (the seedling cohort), only a few will reach maturity and set seed. 3. The diagram also allows for vegetative reproduction, shown as the vegetative daughter connected to the parent plant, (these are called genets). 4. The mature plants will produce seeds, which in turn must pass through the J.L. Harper 1977 environmental filter.
Ecosystem Ecology
Ecosystem ecologist are interested in the response of whole ecosystems.
This diagram shows the mean daily net ecosystem CO2 flux through a simulated wet tundra growing season at 8 C and two atmosphereic CO2 concentration (400 ppm and 800 ppm) and two water table levels (surface and -10 cm).
Billings, W.D. Luken, J.O. Mortensen, D.A. and Peterson, K.M. 1983. Increasing atmospheric carbon dioxide: Possible effects on arctic tundra Oecologia 58: 286-289.
Ecosystem modeling
In the mid 1960s, with the proliferation of computers, ecosystem modeling became a central component of plant ecology. One of the most influential early ecosystem modelers was E.P. Odum, who, based on the ecosystem theories of Tansley, linked plants and animals together through diagrams and computer models of energy-flow, nutrient cycling, and food chains. These types of early models led to very complex models of ecosystems. Models have become increasingly important in all fields of plant ecology. Today they are a key research tool to examine the consequences of global-change. This diagram shows the bioenergetic structure of the detritus-based trophic system in the coastal tundra at Barrow, Alaska.
From MacLean, S.F. 1978. The detritus-based trophic system. In Brown, J. et al. An Arctic Ecosystem: The Coastal Tundra at Barrow, Alaska. Stroudsburg: Dowden, Hutchinson and Ross. Pp. 411-457.
International Biological Programme (IBP)
The concern over the environment in the late 1960s and 1970s led to the development of a large international program called the International Biological Programme to study all the Earths major ecosystems. The earth was divided into biomes, including tundra, grasslands, boreal forests, deserts, and marine seabeds. The IBP formally ended in 1974, and resulted a large number of major books describing the research, such as this one published for the Tundra Biome site at Barrow, Alaska.
Big Ecosystem Science Programs in the Arctic
Long-Term Ecological Research Programme (LTER) (in Alaska there are two LTER sites at Bonanza Creek and Toolik Lake) Research in Arctic Terrestrial Ecosystems (RATE) Response, Resistance, Resilience , and Recovery of Arctic Systems from Disturbance (R4D) FLUX project of the NSF Arctic System Science (ARCSS) program Arctic Transitions in the Land-Atmosphere System (ATLAS)
Composite of covers of Chapin et al., Oechel et al., Reynolds and Tenhunen, Wielgolawski, Tieszen, and Crawford
Flood of Plant Ecology Literature!
My lineage
John Marr Plant ecology teacher, University of Colorado
Patrick J. Webber
Ph.D. Advisor, U. of Colorado Recently retired from Michigan State U. Thesis research on gradient analysis of vegetation of Baffin Island, Canada.
Roland E. Beschel
1928-1971 Plant Ecologist, Canada Sudent of Gams, who helped develop the Northern European School of vegetation science. Pat Webber was his only student. Beschel committed suicide in 1971. He was a prolific writer who is perhaps most famous for his work on using lichens to date the ages glacial surfaces.
Kaye Everett
Ohio State University soil scientists. This man probably had the biggest influence on my field work. Worked together mapping soils and vegetation, and exmining the linkages between arctic soils and vegetation patterns. I spent 15 summers in the field with Kaye. Kaye spent over 30 years in the Arctic and Antarctic.
Vera Komrkov Czecholslovakia
Plant ecologist. Student of Pat Webbers but she was a mature scientist before coming to the University of Colorado. Introduced me to European methods of phytosociology. Led first Womens Expedition to Annapurna, and climbed several peaks in the Himalayas over 8000 m. Mostly she influeneced me by her example of extremely hard work in the field and her European training in a wide range of disciplines that are needed to do plant ecological research.
Fred Danils Phytosociologist, Germany
Co-author of the Circumpolar Arctic Vegetation Map. Extensive vegetation studies in Greenland. Expertise in lichens.
Arve Elvebakk
Phytosociologist from Troms, Norway. Also coauthor of the CAVM.
Some of my heros (not necessarily plant ecologists)
Vera Alexandrova, Russia, geobotanist Carl Benson, UAF, snow scientist and geolgist Dwight Billings, Duke U., plant ecologist, ecophysiology Pete Birkeland, University of Colorado, soil scientist Braun-Blanquet, Switzerland, plant sociologist John Cantlon, Michigan State University, plant ecologist Fred Danils, Germany, phytosociologist Kaye Everett, Ohio State University, soil scientist Heinz Ellenberg, Switzerland, plant ecologist Vera Komrkov, Czechoslovakia, plant ecologist Nadya Mateveyeva, Russia, phytosociologist Pat Webber, Michigan State University, plant ecologist Robert Whittaker, Cornell University, plant ecologist Boris Yurtsev, Russia, geobotanist Fridtjof Nansen, Norwegian Arctic explorer and scientist
Papers for Friday
Cowles, H.C. 1899. The ecological relations of vegetation on the sand dunes of Lake Michigan. Botanical Gazette, Vol. 27.
Billings, W.D. 1952. The environmental complex in relation to plant growth and distribution. Quarterly Review of Biology, 27(3):251265.
PDF versions of the papers are available via the syllabus at http://www.geobotany.uaf.edu/teaching/biol474/474syllabus.html.
Literature Reviews
In short reviews of the literature, please follow these guidelines: 1. Give a short overview of the contents of the paper. Use figures and tables from the paper to help summarize the authors main points. Take no more than 20 minutes for this summary. 2. Give a brief discussion of the relevance of the paper. Why do you feel that this paper is important? Or not important? What was its contribution to the development of plant ecology? Is it still worth reading this paper? What insights has it given you regarding plant ecology? 3. Allow 10 minutes for open discussion and questions at the end.