Abstract
We argue that broad, simple generalizations, not specifically linked to contingencies, will rarely approach truth in ecology and evolutionary biology. This is because most interesting phenomena have multiple, interacting causes. Instead of looking for single universal theories to explain the great diversity of natural systems, we suggest that it would be profitable to develop general explanatory ‘frameworks’. A framework should clearly specify focal levels. The process or pattern that we wish to study defines our level of focus. The set of potential and actual states at the focal level interacts with conditions at the contiguous lower and upper levels of organization, through sets of many-to-one and one-to-many connections. The number of initiating conditions and their permutations at the lower level define the potential states at the focal level, whereas the actual state is constrained by the upper-level boundary conditions. The most useful generalizations are explanatory frameworks, which are road maps to solutions, rather than solutions themselves. Such frameworks outline what is understood about boundary conditions and initiating conditions so that an investigator can pick and choose what is required to effectively understand a specific event or situation. We discuss these relationships in terms of examples involving sex ratio and mating behavior, competitive hierarchies, insect life-histories and the evolution of sex.
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References
Andersen, N. Møller.: 1982, The Semiaquatic Bugs (Hemiptera: Gerromorpha): Phylogeny, Adaptations, Biogeography and Classification, Scandinavian Science Press Ltd, Klampenborg, Denmark.
Allen, T.F.H. and Starr, T.B.: 1982, Hierarchy, Perspectives for Ecological Complexity, University of Chicago Press, Chicago.
Arnqvist, G.: 1988, ‘Mate Guarding and Sperm Displacement in the Water Strider Gerris lateralis Schumm. (Heteroptera: Gerridae)’, Freshwater Biology 19, 269–274.
Arnqvist, G.: 1992a, ‘Precopulatory Fighting in a Water Strider: Inter-sexual Conflict or Mate Assessment’, Animal Behaviour 43, 559–567.
Arnqvist, G.: 1992b, ‘The Effects of Operational Sex Ratio on the Relative Mating Success of Extreme Male Phenotypes in the Water Strider Gerris odontogaster (Zett.) (Heteroptera; Gerridae)’, Animal Behaviour 38, 681–683.
Arnqvist, G.: 1997, ‘The Evolution of Water Strider Mating Systems: Causes and Consequences of Sexual Conflicts’, in J.C. Choe and B.J. Crespi (eds.), The Evolution of Mating Systems in Insects and Arachnids, Cambridge University Press, Cambridge, pp. 146–163.
Bell, G.: 1982, The Masterpiece of Nature, Croom Helm, London.
Bonner, J.T.: 1969, The Scale of Nature, World’s Work Ltd, Kingswood, UK.
Buss, L.: 1987, The Evolution of Individuality, Princeton University Press, Princeton, NJ.
Cole, L.C.: 1954, ‘The Population Consequences of Life-history Phenomena’, Quarterly Review of Biology 29, 103–137.
Eldredge, N.: 1985, Unfinished Synthesis, Oxford University Press, New York.
Greenslade, P.J.M.: 1983, ‘Adversity Selection and the Habitat Templet’, American Naturalist 122, 352–365.
Haila, Y.: 1988, ‘The Multiple Faces of Ecological Theory and Data’, Oikos 53, 408–411.
Haila, Y.: 1990, ‘Toward an Ecological Definition of an Island: a Northwest European Perspective’, Journal of Biogeography 17, 561–568.
Haila, Y. and Levins, R.: 1992, Humanity and Nature - Ecology, Science and Society, Pluto Press, London.
Hairston, N.G. Sr.: 1989, Ecological Experiments, Purpose, Design and Execution, Cambridge University Press, Cambridge.
Jablonski, P. and Vepsäläinen, K.: 1995, ‘Conflict between Sexes in the Water Strider, Gerris lacustris: A Test of Two Hypotheses for Male Guarding Behavior’, Behavioral Ecology 6, 388–392.
Janzen, D.: 1977, ‘What are Dandelions and Aphids?’, American Naturalist 111, 586–589.
Klingenberg, C.P. and Spence, J.R.: 1997, ‘On the Role of Body Size for Life-history Evolution’, Ecological Entomology 22, 55–68.
Kuhn, T.: 1962, The Structure of Scientific Revolutions, University of Chicago Press, Chicago.
Lenton, T.M.: 1998, ‘Gaia and Natural Selection’, Nature 394, 439–447.
Lerner, I.M.: 1954, Genetic Homeostasis, Dover Publications, New York.
Levins, R.: 1966, ‘The Strategy of Model Building in Population Biology’, American Scientist 54, 421–431.
Levins, R.: 1968, Evolution in Changing Environments, Princeton University Press, Princeton, NJ.
Levins, R.: 1974, ‘The Qualitative Analysis of Partially Specified Systems’, Annals of the New York Academy of Sciences 231, 123–138.
Levins, R. and Lewontin, R.: 1985, The Dialectical Biologist, Harvard University Press, Cambridge, MA.
Liem, K.F.: 1990, ‘Key Evolutionary Innovations, Differential Diversity, and Symecomorphis’, in M.H. Nitecki (ed.), Evolutionary Innovations, University of Chicago Press, Chicago, pp. 147–170.
Lovelock, J.E.: 1979, Gaia, a New Look at Life on Earth, Oxford Univeristy Press, Oxford.
Lovelock, J.E.: 1991, Healing Gaia, Practical Medicine for the Planet, Harmony Books, New York.
MacArthur, R.H. and Wilson, E.O.: 1967, The Theory of Island Biogeography, Princeton University Press, Princeton, NJ.
McNaughton, S.J.: 1975, ‘r-& K-selection in Typha’, American Naturalist 109, 251–261.
Maynard Smith, J.: 1978, The Evolution of Sex, Cambridge University Press, Cambridge.
Nummelin, M., Vepsäläinen, K. and Spence, J.R.: 1988, ‘Infection of Gerrid Eggs (Heteroptera: Gerridae) by the Parasitoid Tiphodytes gerriphagus Marchal (Hymenoptera: Scelionidae) in Finland’, Annales Zoologici Fennici 25, 299–302.
O’Neill, R.V., De Angelis, D.L., Waide, J.B. and Allen, T.H.F.: 1986, A Hierarchical Concept of Ecosystems, Princeton University Press, Princeton, NJ.
Peters, R.H.: 1976, ‘Tautology in Evolution and Ecology’, American Naturalist 110, 1–12.
Pianka, E.R.: 1970, ‘On r and K Selection’, American Naturalist 104, 592–597.
Pianka, E.R.: 1972, ‘r and K Selection or b and d Selection’, American Naturalist 106, 581–588.
Popper, K.R.: 1965, Conjectures and Refutations: the Growth of Scientific Knowledge, Harper Torchbooks, New York.
Puccia, C.J. and Levins, R.: 1986, Qualitative Modeling of Complex Biological Systems: Introduction to Loop Analysis and Time Averaging, Harvard University Press, Cambridge, MA.
Roff, D.A.: 1992, The Evolution of Life Histories: Theory and Analysis, Chapman and Hall, New York.
Rowe, L.: 1992, ‘Convenience Polyandry in a Water Strider: Foraging Conflicts and Female Control of Copulation Frequency and Guarding Duration’, Animal Behaviour 44, 189–202.
Rowe, L., Arnqvist, G., Sih, A. and Krupa, J.J.: 1994, ‘Sexual Conflict and the Evolutionary Ecology of Mating Patterns: Water Striders as a Model System’, Trends in Ecology and Evolution 9, 289–293.
Rubenstein, D.I.: 1989, ‘Sperm Competition in the Water Strider, Gerris remigis’, Animal Behaviour 38, 631–636.
Salthe, S.N.: 1985, Evolving Hierarchical Systems, Columbia University Press, New York.
Satchell, J.E.: 1980, ‘r Worms and K Worms: A Basis for Classifying Lumbricid Earthworm Strategies’, in D.L. Dindal (ed.), Soil Biology as Related to Land Use Practices. Proceedings of 7th International Colloquium of Soil Zoology, EPA, Washington, pp. 848–854.
Savolainen, R.: 1990, ‘Colony Success of the Submissive Ant Formica fusca within Territories of the Dominant Formica polyctena’, Ecological Entomology 15, 79–85.
Savolainen, R.: 1991, ‘Interference by Wood Ant Influences Size Selection and Retrieval Rate of Prey by Formica fusca’, Behavioral Ecology and Sociobiology 28, 1–7.
Savolainen, R. and Vepsäläinen, K.: 1988, ‘A Competition Hierarchy Among Boreal Ants: Impact on Resource Partitioning and Community Structure’, Oikos 51, 135–155.
Savolainen, R. and Vepsäläinen, K.: 1989, ‘Niche Differentiation of Ant Species within Territories of the Wood Ant Formica polyctena’, Oikos 56, 3–16.
Savolainen, R., Vepsäläinen, K. and Wuorenrinne, H.: 1989, ‘Ant Assemblages in the Taiga Biome: Testing the Role of Territorial Wood Ants’, Oecologia 81, 481–486.
Shmal’gauzen, I.I.: 1938, Organizm kak Tseloe v Individual’nom i Istoricheskom Razvitii, Izdatel’stvo Akademii Nauk SSSR, Moskva. (in Russian)
Sih, A.: 1988, ‘The Effects of Predators on Habitat Use, Activity and Mating Behavior of a Semi-aquatic Bug’, Animal Behaviour 36, 1846–1848.
Sih, A., Krupa, J. and Travers, S.: 1990, ‘An Experimental Study on the Effects of Predation Risk and Feeding Regime on the Mating Behavior of the Water Strider’, American Nat Uralist 135, 284–290.
Southwood, T.R.E.: 1977, ‘Habitat, the Templet for Ecological Strategies?’, Journal of Animal Ecology 46, 337–365.
Southwood, T.R.E.: 1988, ‘Tactics, Strategies and Templets’, Oikos 52, 3–18.
Spence, J.R.: 1986a, ‘Interactions Between a Scelionid Egg Parasitoid Tiphodytes gerriphagus (Hymenoptera) and its Gerrid Hosts (Heteroptera)’, Canadian Journal of Zoology 64, 2728–2738.
Spence, J.R.: 1986b, ‘Relative impacts of mortality factors in field populations of Gerris buenoi Kirkaldy (Heteroptera: Gerridae)’, Oecologia 70, 68–76.
Spence, J.R.: 1989, ‘The Habitat Templet and Life History Strategies of Pond Skaters (Heteroptera: Gerridae): Reproductive Potential, Phenology and Wing Dimorphism’, Canadian Journal of Zoology 67, 2432–2447.
Spence, J.R. and Andersen, N.M.: 1994, ‘Biology of Waterstriders: Interactions Between Systematics and Ecology’, Annual Review of Entomology 39, 101–128.
Spence, J.R. and Scudder, G.G.E.: 1980, ‘Habitats, Life Cycles and Guild Structure of Waterstriders (Heteroptera: Gerridae) on the Fraser Plateau of British Columbia’, Canadian Entomologist 112, 779–792.
Stearns, S.C.: 1976, ‘Life-history Tactics: A Review of the Ideas’, Quarterly Review of Biology 51, 3–47.
Stearns, S.C.: 1992, The Evolution of Life Histories, Oxford University Press, Oxford.
Taylor, P.: 1989, ‘Revising Models and Generating Theory’, Oikos 54, 121–126.
Thornhill, R. and Alcock, J.: 1983, The Evolution of Insect Mating Systems, Harvard University Press, Cambridge, MA.
Vepsäläinen, K.: 1974, ‘The Wing Lengths, Reproductive Stages and Habitats of Hungarian Gerris Fabr. Species (Heteroptera, Gerridae)’, Annales Academiae Scientiarum Fennicae, Series A IV, Biologica 202, 1–18.
Vepsäläinen, K.: 1978, ‘Wing Dimorphism and Diapause in Gerris: Determination and Adaptive Significance’, in H. Dingle (ed.), The Evolution of Insect Migration and Diapause, Springer-Verlag, New York, pp. 218–253.
Vepsäläinen, K. and Järvinen, O.: 1979, ‘Apomictic Parthenogenesis and the Pattern of the Environment’, American Zoologist 19, 739–751.
Vepsäläinen, K. and Patama, T.: 1983, ‘Allocation of Reproductive Energy in Relation to Pattern of Environment in Five Gerris species’, in V.K. Brown and I. Hodek (eds.), Life Cycle Strategies, Dr. W. Junk bv Publ., The Hague, pp. 189–207.
Vepsäläinen, K. and Pisarski, B.: 1982, ‘Assembly of Island Ant Communities’, Annales Zoologici Fennici 19, 327–335.
Vepsäläinen, K. and Savolainen, R.: 1990, ‘The Effect of Interference by Formicine Ants on the Foraging of Myrmica’, Journal of Animal Ecology 59, 643–654.
Vepsäläinen, K. and Savolainen, R.: 1995, Operational Sex Ratios and Mating Conflict between the Sexes in the Water Strider Gerris lacustris’, American Naturalist 146, 869–880.
Weizenbaum, J.: 1976, Computer Power and Human Reason: From Judgement to Calculation, W. H. Freeman and Company, San Francisco.
Yamamura, N.: 1987, ‘Evolutionarily Stable Strategy (ESS) Models of Copulation and Mate Guarding’, in Y. Itô, J.L. Brown and J. Kikkawa (eds.), Animal Societies: Theories and Facts, Japan Science Society, Tokyo, pp. 197–211.
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Vepsäläinen, K., Spence, J.R. Generalization in Ecology and Evolutionary Biology: From Hypothesis to Paradigm. Biology & Philosophy 15, 211–238 (2000). https://doi.org/10.1023/A:1006636918716
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DOI: https://doi.org/10.1023/A:1006636918716