Download G. Evelyn Hutchinson

G. Evelyn Hutchinson
(1903-1991)
nceas.ucsb.edu
By Angélica Hernández Palma
BIOL 7083
Spring 2014
George Evelyn Hutchinson
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Born in Cambridge, England
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Attended university there
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Began teaching at 23,
in South Africa
Moved to Yale in 1928
Stayed there for 40+
years
Yale University Archives
George Evelyn Hutchinson
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Father of modern ecology
Father of American limnology
Various research interests:
 Limnology
 Biogeochemistry
 Ecosystem,
Phyllis Crowley
community,
population ecology
 Theoretical ecology
“…driven by a fundamental curiosity for nature, coupled with a desire to enhance
appreciation of the diversity of all organisms, even the most cryptic and uncharismatic”
(Gonzalez & Beisner, 2011)
Publications
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First paper at 15! (“A swimming grasshopper”)
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Several important contributions to limnology and community ecology
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limnology.org
“A Treatise on Limnology” 4 volumes (1957 1993)
The Ecological Theater and the Evolutionary Play
(1965)
An Introduction to Population Ecology (1978)
G. Evelyn Hutchinson and the Invention of Modern
Ecology (by N. G. Slack) (2010)
The Art of Ecology: Writings of G. Evelyn
Hutchinson (by Skelly D. K., D. M. Post, & M. D.
Smith – eds.) (2011)
Hutchinson, G.E. 1918. A swimming grasshopper. - Entomological Record and Journal of Variation. 30:138.
Hutchinson, G.E. 1928. The branchial gland of the Cephalopoda- a possible endocrine organ. Nature. 121:674-675.
Hutchinson, G.E., G.E. Pickford, and J.F.M. Schuurman. 1932. A contribution to the hydrobiology of pans and other inland waters of South Africa. Arch. Hydrobiol. 24:1-154.
Hutchinson, G.E. 1932. Experimental studies in ecology. I. The magnesium tolerance of Daphnüdae and its ecological significance. Int. Rev. ges. Hydrobiol. 28:90-108.
Hutchinson, G.E. 1936. The clear mirror. A pattern of life in Goa and in Indian Tibet. Cambridge Univ. Press. 191p.
Hutchinson, G.E. 1937 a. Limnological studies in Indian Tibet. Int. Rev, ges. Hydrobiol. 35:134-177.
Hutchinson, G.E. 1937 b. A contribution to the limnology of arid regions. Trans. Conn. Acad. Arts Sci. 33:47-132.
Hutchinson, G.E. 1938 a. Chemical stratification and lake morphology. Proc. Nat. Acad. Sci. USA. 24:63-69.
Hutchinson, G.E. 1938 b. On the relation between the oxygen deficit and the productivity and typology of lakes. Int. Rev. ges. Hydrobiol. 36:336-355.
Hutchinson, G.E. 1939. Ecological observations on the fishes of Kashmir and Indian Tibet. Ecol. Monogr. 9:142-182.
Hutchinson, G.E., E.S. Deevey and A. Wollack. 1939. The oxidation-reduction potentials of lake waters and their ecological significance. Proc. Nat. Acad. Sci. US. 25:87-90.
Hutchinson, G.E., and A. Wollack. 1940. Studies on Connecticut lake sediments. II. Chemical analyses of a core from Linsley Pond, North Branford. Amer. J. Sci. 238:493-517.
Hutchinson, G.E. 1941. Limnological studies in Connecticut. IV. Mechanism of intermediary metabolism in stratified lakes. Ecol. Monogr. 11:21-60.
Hutchinson, G.E. 1943 a. Marginalia. American Scientist. 31:270.
Hutchinson, G.E. 1943 b. Thiamin in lake waters and aquatic organisms. Arch. Biochem. 2:143-150.
Hutchinson, G.E. 1944. Limnological studies in Connecticut. VII. A critical examination of the supposed relationship between phytoplankton periodicity and chemical changes in lake waters. Ecology.
25:3-26.
Hutchinson, G.E., and J.K. Setlow. 1946. Limnological studies in Connecticut. VIII. The niacin cycle in a small inland lake. Ecology. 27:13-22.
Hutchinson, G.E., and V.T. Bowen. 1947. A direct demonstration of the phosphorus cycle in a small lake. Proc. Nat. Acad. Sci. US. 33:148-153.
Hutchinson, G.E. 1950. Limnological studies of Connecticut. IX. A quantitative radio-chemical study of the phosphorus cycle in Linsley Pond. Ecology. 31:194-203.
Hutchinson, G.E. 1951. Copepodology for the ornithologist. Ecology. 32:571-577.
Hutchinson, G.E. 1953 a. The concept of pattern in ecology. Proc. Acad. Natur. Sci. Phila. 105:1-12.
Hutchinson, G.E. 1953 b. The itinerant ivory tower. Yale University Press.
Hutchinson, G.E., R. Patrick, and E.S. Deevey. 1956. Sediments of Lake Patzcuaro, Michoacan, Mexico. Bull. Geol. Soc. Amer. 67:1491-1504.
Hutchinson, G.E. 1957 a. A treatise on limnology, v. 1. Geography, Physics and Chemistry. Wiley. 1015p.
Hutchinson, G.E. 1957 b. Concluding remarks- Cold Spring Harbor Symposia on Quantitative Biology. 22:415-427. Reprinted in: Classics in Theoretical Biology. Bull. of Math. Biol. 53:193-213.
Hutchinson, G.E. 1959 a. Homage to Santa Rosalia or Why are there so many kinds of animals? Amer. Nat. 93:145-159.
Hutchinson, G.E. 1959 b. Il concetto moderno di niccia ecologica. Mem. Ist. Ital. Idrobiol. 11:9-22.
Hutchinson, G.E. 1961. The paradox of the plankton. Amer. Nat. 95:137-140.
Hutchinson, G.E. 1962. The enchanted voyage and other studies. Yale University Press.
Hutchinson, G.E., and U.M. Cowgill. 1963. Chemical examination of a core from Lake Zeribar, Iran. Science. 140:67-69.
Hutchinson, G.E. 1965. The ecological theater and the evolutionary play. Yale University Press. 139p.
Hutchinson, G.E. 1967. A treatise on limnology, v. 2. Introduction to lake biology and the limnoplankton. Wiley. 1048p.
Hutchinson, G.E. 1970. (ed.) Ianula: An account of the history and development of the Lago di Monterosi, Latium, Italy. Trans. Amer. Philos. Soc. 60(4):178p.
Botkin, D.B., P.A. Jordan, A.S. Dominski, H.S. Lowendorf, and G.E. Hutchinson. 1973. Sodium dynamics in a northern ecosystem. Proc. Nat. Acad. Sci. USA. 70:2745-2748.
Hutchinson, G.E. 1975. A treatise on limnology, v. 3. Limnological Botany. Wiley. 660p.
Hutchinson, G.E. 1978. An introduction to population ecology. Yale University Press.
Hutchinson, G.E. 1979. The kindly fruits of the earth. Recollections of an embryo ecologist. Yale University Press.
Hutchinson, G.E. 1987 a. The ecological niche. Physiology and Ecology Japan. 24:s03-s07.
Hutchinson, G.E. 1987 b. Keep walking- the lecture for the Kyoto Prize 1986. Physiology and Ecology Japan. 24:s81-s87.
Hutchinson, G.E. 1993. A treatise on limnology, v. 4. The Zoobenthos. Wiley. 964p.
Awards
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Many honorary doctorates (one from Cambridge
University)
Benjamin Franklin Medal (1979)
for “developing scientific basis of ecology”
Kyoto Prize in Basic Science (1986)
National Medal of Science (1991 posthumous)
Yale University Library
 Leidy Medal, Philadelphia Academy of Natural Sciences (1955)
 Naumann Medal, International Association of Theoretical and Applied Limnology
(1959)
 Eminent Ecologist Award, Ecological Society of America (1962)
 Tyler Award (1974)
 Frederick Garner Cottrell Award for Environmental Quality, National Academy of
Sciences, USA (1974)
 Daniel Giraud Elliot Medal, National Academy of Sciences (1984)
HUTCHINSON’S INTELLECTUAL FAMILY TREE
From Edmondson (1971)
Concluding Remarks
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Hutchinson, G.E. (1957). Concluding remarksCold Spring Harbor Symposia on Quantitative
Biology. 22:415-427
Cited >2000 times
Establishment of the “Hutchinsonian niche”
concept
Concluding Remarks
Humidity (X2)
X1, X2, …Xn  independent
environmental variables
Ecological factors relative to S1
Temperature (X1)
Humidity (X2)
n-dimensional hypervolume 
state of environment that permits
S1 to exist indefinitely
Temperature (X1)
FUNDAMENTAL NICHE
(defines a species’ ecological properties)
Concluding Remarks
No
persistence
Equal probability of
persistence
Erik Pianka
Concluding Remarks
REALIZED NICHE
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
Erik Pianka
Fundamental niche
never fully realized
(due to interspecific
interactions –
competition, predation)
Actual fraction of
fundamental niche that
a species realizes
Concluding Remarks
Included:
1. S2 is superior (dashed curve), persists and S1
reduces utilization of shared resources
2. S1 is superior (solid curve), S2 excluded and S1
uses entire resource
Refuge
Equal overlap: competition is equal and opposite
Unequal overlap: competition is not equal and
opposite
Superior
competitor
gets this
Abutting: no direct competition. May indicate an
avoidance of it
Disjunct: no competition. Completely different
fundamental niches
Erik Pianka
antique-prints.de
Shrine of
Santa Rosalia
near Palermo,
Sicily
Homage to Santa Rosalia
or Why are there so many kinds of animals?
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Water-bugs (Corixa)
C. punctata  larger. Only females. Ending breeding
C. affinis  smaller. Equal number of both sexes.
Starting breeding
C. punctata wikipedia.org C. affinis
bugguide.net
Federico Marrone
The pond of Santa Rosalia on Monte Pellegrino
(Palermo, Italy)
Homage to Santa Rosalia
or Why are there so many kinds of animals?
Why larger species should breed first?
Why only 2 species and not
20 or 200?
Why there are such an
enormous number of animal
species?
Homage to Santa Rosalia
or Why are there so many kinds of animals?
INTERRELATIONS OF FOOD CHAINS
 links in food web   stability in the community
(MacArthur) evolution of communities:
  efficient spp replace   efficient spp
  stable comms replace   stable comms
Entry of new species:
1. Completely displace an old species
(no change in stability)
2. Occupy an unfilled niche
(provide new links; increase stability)
3. Partition a niche with pre-existing species
biologycorner.com
Homage to Santa Rosalia
or Why are there so many kinds of animals?
Early in a community many niches empty  invasion easier
Invaders:
 Oust a species
 Compete for marginal parts of niche
Entry of invader 
Most likely when
a species is
fluctuating and is
underrepresented
 pop of orig species +  stability by reducing fluctuations
 Loss of niche space compensated by reduction in
amplitude of fluctuations
Diverse communities better able to persist!!
Homage to Santa Rosalia
or Why are there so many kinds of animals?
EFFECTS OF TERRESTRIAL PLANTS
 What determines the number of food chains in a
community? Diversity of primary producers
Major source of terrestrial diversity introduced by evolution
of ~ 200.000 species or flowering plants
~ 750.000 species of insects
Why are there so many kinds of
plants!?
Limits to diversity…
Homage to Santa Rosalia
or Why are there so many kinds of animals?
FOOD CHAINS
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Elton (1927) the predator at each level is
larger than its prey
Each predator 2X size  5th animal’s
population 10-4 of the 1st
20%
anselm.edu
Eltonian food-chain
cannot give any
great diversity!
wikipedia.org
Homage to Santa Rosalia
or Why are there so many kinds of animals?
FOOD CHAINS – NATURAL SELECTION
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Selective force operating on
food chains  may limit
diversity
Natural selection  maintain
efficiency of transfer at a
maximum
Shortening of
food chains!
 predatory
efficiency (n)
 extinction
risk (n-1)
Adapts to eat
(n-2) or extinct
Lengthening  development of a
new carnivore link (empty niche)
Not likely to be easy…
wikipedia.org
Homage to Santa Rosalia
or Why are there so many kinds of animals?
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Total biomass (primary
productivity)  short
growing season
Small populations (rarer
species so rare they do
not exist at all)
Limited number of niches
environment.nationalgeographic.com

Size of habitat available
for colonization (islands)
Homage to Santa Rosalia
or Why are there so many kinds of animals?
NICHE REQUIREMENTS
How much difference between two species at the same
level is needed to prevent them from occupying the
same niche?
Character displacement (Brown & Wilson 1956)
divergence when two allopatric species of comparable
niche become sympatric
Kind of difference necessary to
permit two species to co-occur in
different niches but at the same
level of a food web
In sympatry
ratio larger : smaller
~ 1.3:1
(“Hutchinson’s ratio”)
• Birds: culmen length
• Mammals: skull size
bioserv.fiu.edu
Homage to Santa Rosalia
or Why are there so many kinds of animals?
MOSAIC NATURE OF THE ENVIRONMENT
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(except for open water)  every area has some local
diversity
Depends largely on size of organisms
(always) more species of
small-medium organisms
than large ones
“Small size permits animals to
become specialized to conditions
offered by small elements of the
environmental mosaic”
300 m
3 km
10 km
Homage to Santa Rosalia
or Why are there so many kinds of animals?
“But perhaps Santa Rosalia would
find at this point that we are
speculating too freely, so for the
moment, while under her patronge, I
will say no more.”
Homage to Santa Rosalia
or Why are there so many kinds of animals?
Role of energy in food chains
 Effects of productivity
 Available habitat
 Community stability
 Environmental grain

MAINTENANCE
OF
BIODIVERSITY
How is it possible for a number of species to
coexist in a relatively unstructured environment
all competing for the same sorts of materials?
Phytoplankton
of large
bodies of
water
pnas.org
The paradox of the plankton
PHYTOPLANKTON…
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Phototrophs able to reproduce and build up populations
in inorganic media with a source of CO2
+ Inorganic N, S, P
+ Na, K, Mg, Ca, Si, Fe, Mn, B, Cl, Cu, Zn, Mo, Co, V
(required in small concentrations – not by all)
Vitamins (thiamin, B12, biotin)
Natural waters in summer very
nutrient deficient  severe
competition
wikipedia.org
The paradox of the plankton
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Principle of competitive exclusion (Gause 1935; Hardin 1960)
One species alone would outcompete all the others
 Based on equilibrium conditions
 Never obtained (environm. var.)
 Little empirical interest
Competition between 2 laboratory
populations of Paramecium.
Gause (1935)
Sylvia S. Mader
The paradox of the plankton
Diversity of phytoplankton maybe due to failure to
achieve equilibrium as external factors changed
tc  time to completely replace a species (reproductive rate)
te  time for a significant change in environment to occur
tc << te  competitive exclusion at equilibrium complete
before the environment changes significantly
tc ≈ te  no equilibrium achieved
tc >> te  competitive exclusion occurring in a changing
environment to the full range of which individual competitors
would have to be adapted to live alone
The paradox of the plankton
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Light gradient in epilimnion?
Chemical conditions at surface film?
Motility?
Small chances for any organism
to remain permanently in a
particular range of intensities
(turbulence, surface winds)
Not many (physical) opportunities
for niche diversification
rmbel.info
The paradox of the plankton
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Under some conditions, commensal/symbiotic
species can occupy the same niche
Phytoplankton some species require vitamins and
others do not
 efficient species requires vitamins
 efficient species produces vitamins
Mixed
equilibrium
populations
The paradox of the plankton
Role of predation…
Prey species A  Predator X (limiting)
Prey species B  Predator X (not limiting) / Predator Y (limiting)
COEXISTENCE OF THE TWO PREY SPECIES
LIKELY OUTCOME
corbisimages.com
“… for the moment I am content that
its use has demonstrated possible
ways of looking at the problem [of
plankton diversity] and, I hope, of
presenting that problem to you”
The paradox of the plankton
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Competitive exclusion can be avoided in nature
Competitive exclusion many never be achieved because of the
rapid change at which the environments change
Even for short time periods, non-equilibrium rather than
equilibrium conditions could be the rule
Different species favored under different sets of
environmental conditions. If enough changes through time, no
single competitor could remain superior long enough to
exclude other species
Role of environment in structuring communities
Shed light on Connell’s “intermediate disturbance hypothesis”
“…he found therein several questions on
some of the most intriguing arguments of
ecology and virtually no answers to the
questions themselves. This left him rather
confused, because, in his image of science,
scientific literature existed to give answers
rather than disseminating doubts.”
Naselli-Flores & Rossetti (2010)
References
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Brown, W. L. & E. 0. Wilson. 1956. Character displacement. Systematic Zoology 5: 49-64.
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Elton, C. S. 1927. Animal Ecology. University of Chicago Press. 209 p.
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Gause, G. F. 1935. Experimental demonstration of Volterra's periodic oscillations in the numbers of animals. Journal of Experimental
Biology. 12:44-48.
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Gonzalez, A. & B. Beisner. 2011. Homage to G. Evelyn. Hutchinson. Book Review. Conservation Biology. 25:1253-1262.
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Hardin, G. 1960. The competitive exclusion principle. Science 131: 1292- 1298.
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Hutchinson, G.E. 1957 a. A treatise on limnology, v. 1. Geography, Physics and Chemistry. Wiley. 1015p.

Hutchinson, G.E. 1957 b. Concluding remarks- Cold Spring Harbor Symposia on Quantitative Biology. 22:415-427. Reprinted in:
Classics in Theoretical Biology. Bull. of Math. Biol. 53:193-213.

Hutchinson, G.E. 1959 a. Homage to Santa Rosalia or Why are there so many kinds of animals? Amer. Nat. 93:145-159.

Hutchinson, G.E. 1961. The paradox of the plankton. Amer. Nat. 95:137-140.

Hutchinson, G.E. 1965. The ecological theater and the evolutionary play. Yale University Press. 139p.

Hutchinson, G.E. 1967. A treatise on limnology, v. 2. Introduction to lake biology and the limnoplankton. Wiley. 1048p

Hutchinson, G.E. 1975. A treatise on limnology, v. 3. Limnological Botany. Wiley. 660p.

Hutchinson, G.E. 1978. An introduction to population ecology. Yale University Press.

Hutchinson, G.E. 1993. A treatise on limnology, v. 4. The Zoobenthos. Wiley. 964p.

Naselli-Flores, L. & G. Rossetti. 2010. Santa Rosalia, the icon of biodiversity. Hydrobiologia. 653:235-243.
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Skelly, D. K., D. M. Post & M. D. Smith (eds.) (2011). The Art of Ecology: Writings of G. Evelyn Hutchinson. Yale University Press. 368 p.

Slack, N. 2010. G. Evelyn Hutchinson and the Invention of Modern Ecology. Yale University Press. 480 p.