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G. Evelyn Hutchinson (1903-1991) nceas.ucsb.edu By Angélica Hernández Palma BIOL 7083 Spring 2014 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, 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 First paper at 15! (“A swimming grasshopper”) Several important contributions to limnology and community ecology 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 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 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 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? 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 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 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? 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 (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… 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 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 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 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 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.