Abstract
Many authors, including paleobiologists, cladists and so on, adopt a nested hierarchical viewpoint to examine the relationships among different levels of biological organization. Furthermore, species are often considered to be unique entities in functioning evolutionary processes and one of the individuals forming a nested hierarchy.
I have attempted to show that such a hierarchical view is inadequate in evolutionary biology. We should define units depending on what we are trying to explain. Units that play an important role in evolution and ecology do not necessarily form a nested hierarchy. Also the relationships among genealogies at different levels are not simply nested. I have attempted to distinguish the different characteristics of passages when they are used for different purposes of explanation. In my analysis, species and monophyletic taxa cannot be uniquely defined as single units that function in ecological and evolutionary processes.
The view discussed in this paper may provide a more general basis for testing competing theories in evolution, and provide new insights for future empirical studies.
Similar content being viewed by others
References
Arthur, W.: 1984, Mechanisms of Morphological Evolution: A Combined Genetic, Developmental and Ecological Approach, J. Wiley & Son, Chichester.
Barton, N. H. and B. Charlesworth: 1984, ‘Genetic Revolutions, Founder Effects and Speciation’, Ann. Rev. Ecol. Syst. 15, 133–164.
Brandon, R. N. and R. M. Burian (eds.): 1984, Genes, Organisms, Populations: Controversies Over the Units of Selection, MIT Press, Cambridge.
Bush, G. L., S. M. Case, A. C. Wilson and J. L. Patton: 1977, ‘Rapid Speciation and Chromosomal Evolution in Mammals’, Proc. Natl. Acad. Sci. USA 74, 3942–3946.
Charlesworth, B., R. Lande and M. Slatkin: 1982, ‘A Neo-Darwinian Commentary on Macroevolution’, Evolution 36, 474–498.
Chalmers, A. F.: 1982, What is This Thing Called Science? 2nd ed., Univ. Queensland Press, Queensland.
Damuth, J.: 1985, ‘Selection Among “Species”: A Formation in Terms of Natural Functional Units’, Evolution 39, 1132–1146.
Dawkins, R.: 1982a, The Extended Phenotype: The Gene as the Unit of Selection, Freeman, Oxford.
Dawkins, R.: 1982b, ‘Replicators and Vehicles’, in King's College Sociobiology Group (ed.), Current Problems in Sociobiology, Cambridge Univ. Press, Cambridge, pp. 45–64.
Dover, G. A. and R. B. Flavell (eds.): 1982, Genome Evolution. Academic Press, London.
Ehrlich, P. R. and P. H. Raven: 1969, ‘Differentiation of Populations’, Science 165, 1228–1232.
Eldredge, N.: 1985, Unfinished Synthesis: Biological Hierarchies and Modern Evolutionary Thought, Oxford Univ. Press, New York.
Eldredge, N. and S. J. Gould: 1972, ‘Punctuated Equilibria: An Alternative to Phyletic Gradualism’, in T. J. M. Schopf (ed.), Models in Paleontology, Freeman, San Francisco, pp. 88–115.
Eldredge, N. and J. Cracraft: 1980, Phylogenetic Patterns and the Evolutionary Process: Methods and Theory in Comparative Biology, Columbia Univ. Press, New York.
Eldredge, N. and S. N. Salthe: 1984, ‘Hierarchy and Evolution’, Oxford Surveys Evol. Biol. 1, 184–208.
Endler, J. A.: 1977, Geographic Variation, Speciation and Clines, Princeton Univ. Press, Princeton.
Ferris, S. D., R. D. Sage, C. M. Huang, J. T. Nielsen, U. Ritte and A. C. Wilson: 1983, ‘Flow of Mitochondrial DNA Across a Species Boundary’, Proc. Natl. Acad. Sci. USA. 80, 2290–2294.
Feyerbend, P. K.: 1975, Against Method, New Left Books, London.
Gallez, G. P. and L. D. Gottlieb: 1982, ‘Genetic Evidence for the Hybrid Origin of the Diploid Plant, Stephanomeria Diegensis’ Evolution 36, 1158–1167.
Ghiselin, M. T.: 1974, ‘A radical Solution to the Species Problem’. Syst. Zool. 23, 536–544.
Gilinsky, N. L.: 1986, ‘Species Selection as a Causal Process’, Evol. Biol. 20, 249–273.
Gould, S. J.: 1980, ‘Is a New and General Theory of Evolution Emerging?’ Paleobiol. 6, 119–130.
Gould, S. J.: 1982, ‘The Meaning of Punctuated Equilibrium and its Role in Validating a Hierarchical Approach to Macroevolution. in R. Milkman (ed.), Perspective on Evolution, Sinauer, Sunderland, pp. 83–104.
Gould, S. J. and N. Eldredge: 1977, ‘Punctuated Equilibria: The Tempo and Mode of Evolution Reconsidered’, Paleobiol. 3, 115–151.
Goodwin, B. C.: 1984, ‘Changing from an Evolutionary to Generative Paradigm in Biology’ in J. W. Pollard (ed.), Evolutionary Theory: Paths into the Future, John Wiley & Sons, Chichester, pp. 99–120.
Green, M. M.: 1980, ‘Transposable Genetic Elements in Drosophila and Other Diptera’, Ann. Rev. Genet. 14, 109–120.
Hanson, N. R.: 1969, Perception and Discovery, Freeman, San Francisco.
Hoffman, A.: 1984, ‘Species Selection’, Evol. Biol. 18, 1–20.
Holsinger, K. E.: 1984, ‘The Nature of Biological Species’, Phil. Sci. 51, 293–307.
Hull, D. L.: 1976, ‘Are species Really Individuals? Syst. Zool. 25, 174–191.
Hull, D. L.: 1980, ‘Individuality and Selection’, Ann. Rev. Ecol. Syst. 11, 311–322.
Hull, D. L.: 1981, ‘Units of Evolution: A Metaphysical Essay’, in J. Jensen and R. Harre (eds.), The Philosophy of Evolution, Harvester Press, Brinhton, pp. 23–44.
Jablonski, D.: 1986, ‘Background and Mass Extinctions: The Alternation of Macroevolutionary Regimes’, Science 231, 129–133.
Kimura, M.: 1983, The Neutral Theory of Molecular Evolution, Cambridge Univ. Press, Cambridge.
Kitcher, P.: 1984, ‘Species’, Phil. Sci. 51, 308–333.
Kuhn, T.: 1970, The Structure of Scientific Revolution, 2nd ed. Univ. Chicago Press, Chicago.
Lakatos, I.: 1978, Imre Lalatos, Philosophical Papers Vol. 1: the Methodology of Scientific Research Programs. (J. Worrel and G. Currie eds.), Cambridge Univ. Press, Cambridge.
Lande, R.: 1980, ‘Microevolution in Relation to Macroevolution’, Paleobiol. 6, 235–238.
Lande, R. and S. J. Arnold: 1983, ‘The Measurement of Selection on Correlated Characters’, Evolution 37, 1210–1226.
LeRoith, D. and J. Roth: 1984, ‘Vertebrate Hormones and Neuropeptides in Microbes: Evolutionary Origin of Intercellular Communication’, Frontiers in Neuroendcrinology 8, 1–25.
Margulis, L.: 1981, Symbiosis in Cell Evolution: Life and Its Environment of the Early Earth, W. H. Freeman, San Francisco.
Maynard Smith, J.: 1983, ‘The Genetics of Stasis and Punctuation’, Ann. Rev. Genet. 17, 11–25.
Mayr, E.: 1963, Animal Species and Evolution, Harvard Univ. Press, Cambridge.
Mishler, B. D.: 1985, ‘The Morphological, Developmental, and Phylogenetic Basis of Species Concepts in Bryophytes’, The Bryologist 88, 207–214.
Mishler, B. D. and M. J. Donoghue: 1982, ‘Species Concepts: A Case for Pluralism’, Syst. Zool. 31, 491–503.
Nei, M.: 1971, ‘Interspecific Gene Differences and Evolutionary Time Estimated from Electrophoretic Data or Protein Identity’, Am. Nat. 105, 385–398.
Ohno, S.: 1970, Evolution by Gene Duplication, Springer-Verlag, New York.
Oster, G. and P. Alberch: 1982, ‘Evolution and Bifurcation of Developmental Programs’, Evolution 36, 444–459.
Paterson, H. E. H.: 1978, ‘More Evidence Against Speciation by Reinforcement’, S. Afr. J. Sci. 74, 369–371.
Paterson, H. E. H.: 1982, ‘Perspective on Speciation by Reinforcement’, S. Afr. J. Sci. 78, 53–57.
Pashley, D. P., K. S. Rai and D. N. Pashley: 1985, ‘Patterns of Allozyme Relationships Compared with Morphology, Hybridization, and Geologic History in Allopatric Island-Dwelling Mosquitoes’, Evolution 39, 985–997.
Powell, J. R.: 1983, ‘Interspecific Cytoplasmic Gene Flow in the Absence of Nuclear Gene Flow Evidence Form Drosophila’, Proc. Natl. Acad. Sci. USA. 80, 482–495.
Ricklefs, R. E.: 1979, Ecology, 2nd ed.. Chiron Press, New York.
Roizes, G. P. and M. Pages: 1982, ‘Conserved and Divergent Sequences of Bovine Satellite DNA’, in G. A. Dover and R. B. Flavell (eds.), Genome Evolution, Academic Press, London, pp. 95–111.
Simpson, G. G.: 1961, Principles of Animal Taxonomy, Columbia Univ. Press, New York.
Slack, J.: 1984, ‘A Rosetta Stone for Pattern Formation in Animals?’ Nature 310, 364–365.
Sober, F.: 1984, The Nature of Selection: Evolutionary Theory in Philosophical Focus, MIT Press, Mass.
Spolsky, C. and T. Uzzel: 1984, ‘Natural Interspecies Transfer Mitochondria DNA Amphibians’, Proc. Natl. Acad. Sci. USA 81, 5802–5805.
Stanley, S. M.: 1979, Macroevolution: Patter and Process, Freeman, San Francisco.
Stebbins, G. G.: 1950, Variation and Evolution in Plants, Columbia Univ. Press, New York.
Takahata, N. and M. Nei: 1986, ‘Gene Genealogy and Variance of Interpopulational Nucleotide Differences’, Genetics 110, 325–344.
Templeton, A. R.: 1981, ‘Mechanisms of Speciation: A Population Genetic Approach’, Ann. Rev. Ecol. Syst. 12, 23–48.
Van Valen, L.: 1976, ‘Ecological Species, Multispecies and Oaks’, Taxon 25, 233–239.
Van Valen, L.: 1982, ‘Integration of Species: Stasis and Biogeography’, Evol. Theor. 6, 99–112.
Vrba, E. and N. Eldredge: 1984, ‘Individuals, Hierarchies and Processes: Towards a More Complete Evolutionary Theory’, Paleobiol. 10, 146–171.
Webster, G.: 1984, ‘The Relations of Natural Forms’, H-W. Ho, and P. T. Saunders (eds.), Beyond Neo-Darwinism, Academic Press, New York, pp. 193–217.
White, M. J. D.: 1987, Mode of Speciation, Freeman, San Francisco.
Wiley, E. O.: 1981, Phylogenetics: The Theory and Practice of Phylogenetic Systematics, Wiley and Sons, New York.
Williams, G. C.: 1985, ‘A Defense of Reductionism in Evolutionary Biology’, Oxford Sur. Evol. Biol. 2, 1–27.
Wilson, D. S.: 1980, The Natural Selection of Populations and Communities, Benjamin/Commings, Menlo Parks.
Wilson, D. S.: 1986, ‘Adaptive Indirect Effect’, in J. Diamond and T. J. Case (eds.), Community Ecology, Harper & Row, New York, pp. 437–444.
Wu, J. C. and L. Manuelidis: 1980, ‘Sequence Definition and Organization of a Human Repeated DNA’, J. Mol. Biol. 142, 386.
Yonekawa, H. et al.: 1982, ‘Origins of Laboratory Mice Deduced from Restriction Patterns of Mitochondrial DNA’, Differentiation 22, 222–226.