G. Evelyn Hutchinson · Hutchinson, G.E. 1918.A swimming grasshopper. - Entomological Record and...

G. Evelyn Hutchinson (1903-1991)

By Angélica Hernández Palma

BIOL 7083

Spring 2014

nceas.ucsb.edu

George Evelyn Hutchinson

Born in Cambridge, England

Attended university there

Began teaching at 23, in South Africa

Moved to Yale in 1928

Stayed there for 40+ years

Yale University Archives

George Evelyn Hutchinson

Father of modern ecology

Father of American limnology

Various research interests:

Limnology

Biogeochemistry

Ecosystem, 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)

Phyllis Crowley

Publications

First paper at 15! (“A swimming grasshopper”)

Several important contributions to limnology and community ecology

“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)

limnology.org

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

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)

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)

Yale University Library

HUTCHINSON’S INTELLECTUAL FAMILY TREE From Edmondson (1971)

Concluding Remarks

Hutchinson, G.E. (1957). Concluding remarks-

Cold Spring Harbor Symposia on Quantitative

Biology. 22:415-427

Cited >2000 times

Establishment of the “Hutchinsonian niche”

concept

X1, X2, …Xn independent

environmental variables

Ecological factors relative to S1

n-dimensional hypervolume

state of environment that permits

S1 to exist indefinitely

FUNDAMENTAL NICHE

Concluding Remarks

Temperature (X1)

Hum

idity (X

2)

Temperature (X1)

Hum

idity (X

2)

(defines a species’ ecological properties)

Concluding Remarks

No

persistence

Equal probability of

persistence

Erik Pianka

Fundamental niche

never fully realized

(due to interspecific

interactions –

competition, predation)

Actual fraction of

fundamental niche that

a species realizes

Concluding Remarks

REALIZED NICHE

Erik Pianka

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

Equal overlap: competition is equal and opposite

Unequal overlap: competition is not equal and

opposite

Refuge

Superior

competitor

gets this

Abutting: no direct competition. May indicate an

avoidance of it

Disjunct: no competition. Completely different

fundamental niches

Erik Pianka

Shrine of

Santa Rosalia

near Palermo,

Sicily antique-prints.de

Homage to Santa Rosalia

or Why are there so many kinds of animals?

Water-bugs (Corixa)

C. punctata larger. Only females. Ending breeding

C. affinis smaller. Equal number of both sexes.

Starting breeding

The pond of Santa Rosalia on Monte Pellegrino

(Palermo, Italy)

C. punctata wikipedia.org C. affinis bugguide.net

Federico Marrone



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?



INTERRELATIONS OF FOOD CHAINS

links in food web stability in the community

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

(MacArthur) evolution of communities:

efficient spp replace efficient spp

stable comms replace stable comms

biologycorner.com



Early in a community many niches empty invasion easier

Invaders:

Oust a species

Compete for marginal parts of niche

Entry of invader 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!!

Most likely when

a species is

fluctuating and is

underrepresented



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…



FOOD CHAINS

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

anselm.edu wikipedia.org

20%

Eltonian food-chain

cannot give any

great diversity!



FOOD CHAINS – NATURAL SELECTION

Selective force operating on food chains may limit diversity

Natural selection maintain efficiency of transfer at a maximum

predatory

efficiency (n)

extinction

risk (n-1)

Adapts to eat

(n-2) or extinct

Shortening of

food chains!

Lengthening development of a

new carnivore link (empty niche)

wikipedia.org

Not likely to be easy…



Total biomass (primary

productivity) short

growing season

Small populations (rarer

species so rare they do

not exist at all)

Limited number of niches

Size of habitat available

for colonization (islands)

environment.nationalgeographic.com



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

In sympatry

ratio larger : smaller

~ 1.3:1

(“Hutchinson’s ratio”)

Kind of difference necessary to

permit two species to co-occur in

different niches but at the same

level of a food web

• Birds: culmen length

• Mammals: skull size

bioserv.fiu.edu



MOSAIC NATURE OF THE ENVIRONMENT

(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

10 km

3 km

300 m “Small size permits animals to

become specialized to conditions

offered by small elements of the

environmental mosaic”



“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.”



Role of energy in food chains

Effects of productivity

Available habitat

Community stability

Environmental grain

MAINTENANCE

OF

BIODIVERSITY

Phytoplankton

of large

bodies of

water

pnas.org

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…

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

The paradox of the plankton

wikipedia.org


Principle of competitive exclusion (Gause 1935; Hardin 1960)

One species alone would outcompete all the others

Competition between 2 laboratory

populations of Paramecium.

Gause (1935)

Sylvia S. Mader

Based on equilibrium conditions

Never obtained (environm. var.)

Little empirical interest


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


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


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


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”


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

Brown, W. L. & E. 0. Wilson. 1956. Character displacement. Systematic Zoology 5: 49-64.

Elton, C. S. 1927. Animal Ecology. University of Chicago Press. 209 p.

Gause, G. F. 1935. Experimental demonstration of Volterra's periodic oscillations in the numbers of animals. Journal of Experimental

Biology. 12:44-48.

Gonzalez, A. & B. Beisner. 2011. Homage to G. Evelyn. Hutchinson. Book Review. Conservation Biology. 25:1253-1262.

Hardin, G. 1960. The competitive exclusion principle. Science 131: 1292- 1298.

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.

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.

G. Evelyn Hutchinson · Hutchinson, G.E. 1918.A swimming grasshopper. - Entomological Record and...

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