Lesson 2 Brief history of plant ecology

Lesson 2
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
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Explored the world’s biomes
seeking to develop a consistent
means to describe the diverse
plant life of the globe.
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Von Humboldt traveled to Central
America,
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Alexander von Humboldt
(1773-1858)
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.
Aimé Bonpland
(1773-1858)
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Von Humboldt’s 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.
Humboldt’s 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_
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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
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J.F. Schouw (1789-1852)
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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.
Anton Kerner von Marilaun (1831-1898) (Kerner)
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Transplant gardens in Tyrolean Alps (300 species). Distinguished inheritable from
environmentally-affected factors. Beautiful vegetation descriptions and understanding of
succession in :
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___________. 1863. Plant life of the Danube Basin. Translated by H.S. Conrad.
1951.
August Grisebach (1814-1879)
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Succeeded Willdenow at Göttingberg. Described more than 50 major vegetation formations
worldwide in modern physiognomic terms, relating their distribution to climatic factors.
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Johannes Eugenius Bülow Warming (1841-1924)
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Andreas Franz Wilhelm Schimper (1856-1911)
Johannes Eugenius
Bülow Warming (18411924)
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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)
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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.
• Raunkiær (1934), The Life
Forms of Plants and
Statistical Plant
Geography.
Raunkiaer’s growth forms
European plant
community
classification phase
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In Europe in the early 20th
Century, most vegetation
scientists were occupied by
classification of vegetation.
Most famous was Josias BruanBlanquet (1884-1980).
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Josias Braun-Blanquet
(1884-1980)
Followed in path of Kerner.
Developed methods of
community sampling, data
reduction, association
nomenclature.
Cooperated with Rübel 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).
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)
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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 1800’s and early 1900’s
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:
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Jozef Paczoski (1864-1941)
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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)
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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)
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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.
Merrriam’s
life zones
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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.
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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
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Pond succession in Ohio based on the work of Dachnowski.
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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
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Perhaps the most famous of the
early American ecologists. Born,
raised, and educated in
Nebraska.
Enchanted with the stories of
Lewis and Clark’s 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.
Clement’s detailed
observations of plants
and plant communities
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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
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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.
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This “organismic approach”
was disputed--rather quietly
by many prominent
ecologists, and rather
heatedly by Henry Gleason.
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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)
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Student of Henry Cowles;
primarily a taxonomist, with
a strong grounding in the
'physiographic ecology' of
Cowles.
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.
Gleason’s “individualistic
hypothesis”
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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.
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Studied the Mississippi Valley and
the changes in community
composition along the length of the
river.
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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)
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University of Wisconsin, professor of
botany.
Plant physiologist, orchid specialist.
Strongly influenced by Gleason’s
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
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Tilia americana
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Bray and Curtis Ordination
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.
Robert H. Whittaker
(1920-1980)
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Probably the most famous
vegetation scientist in the US.
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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).
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Produced 5-kingdom
classification for organisms.
Gradient analysis of Smokey Mountain ecosystems
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Like Curtis, Whittaker,
developed methods to portray
vegetation along complex
environmental gradients.
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Primarily examined vegetation
along elevation gradients in
mountain ecosystems (Smokey
Mountains, the Siskiyou
Mountains of Oregon, and the
Santa Cantalina Mountains in
Arizona).
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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. Müller-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
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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 1940’s and 1950’s 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
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Ecosystem ecologist are
interested in the response of
whole ecosystems.
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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
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.
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In the mid 1960’s, with the
proliferation of computers,
ecosystem modeling became a
central component of plant ecology.
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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.
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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.
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This diagram shows the bioenergetic
structure of the detritus-based
trophic system in the coastal tundra
at Barrow, Alaska.
International Biological
Programme (IBP)
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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 Earth’s
major ecosystems.
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The earth was divided into biomes,
including tundra, grasslands,
boreal forests, deserts, and marine
seabeds.
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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 Komárková
Czecholslovakia
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Plant ecologist.
Student of Pat Webber’s but she was a
mature scientist before coming to the
University of Colorado.
Introduced me to European methods of
phytosociology.
Led first Women’s 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 Daniëls
Phytosociologist, Germany
• Co-author of
the
Circumpolar
Arctic
Vegetation
Map.
• Extensive
vegetation
studies in
Greenland.
• Expertise in
lichens.
Arve Elvebakk
• Phytosociologist
from Tromsø,
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 Daniëls, Germany, phytosociologist
• Kaye Everett, Ohio State University, soil scientist
• Heinz Ellenberg, Switzerland, plant ecologist
• Vera Komárková, 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.