Key Scientists Advocating non-Gradualist Evolution    
in the 19th and 20th Centuries


Non-gradual Evolutionary Process


William Bateson (1861-1926)

Discontinuous variation

(Bateson 1894; Gillham 2001; Gillham 2001; Bateson 2002; Peterson 2008)

Hugo de Vries (1848-1935)

Abrupt mutational variation

(de Vries 1901-1903; de Vries 1905)

Konstantin Mereschkowsky  (1855–1921)

Evolution by symbiogenesis

(Mereschkowsky 1910; Mereschkowsky 1926)

Ivan E. Wallin (1883-1969)

Evolution by symbiogenesis (“Symbionticism”)

(Wallin 1922; Wallin 1922; Wallin 1927)

Kozo-Polyansky, Boris Mikhailovich  (1890-1957)

Evolution by symbiogenesis

(Kozo-Polyansky 1924 (2010, English translation by Victor Fet))

George Gaylord Simpson (1902-1984)

Quantum evolution

(Simpson G.G 1944; Gould 1980)

Richard Goldschmidt (1878-1958)

“Hopeful monsters” formed by redirecting developmental programs

(Goldschmidt 1940; Gould 1977; Dietrich 2003)

George Ledyard Stebbins (1906-2000)

Hybrid Speciation (“Cataclysmic Evolution”)

(Stebbins 1940; Stebbins 1951; Stebbins 1971; Stebbins and Ayala 1981)

Niles Eldredge  (b. 1943) and Stephen J. Gould (1941-2002)

Punctuated equilibrium

(Eldredge and Gould 1972; Gould 1983; Gould and Eldredge 1993)

Lynn Margulis (1938-2011)

Evolution by symbiogenesis

(Sagan 1967; Margulis 1970; Margulis 1971; Margulis 1981; Sapp 1994; Margulis and Sagan 2002; Sapp 2010)





Bateson, P. (2002). "William Bateson: a biologist ahead of his time." J Genet 81(2): 49-58.

Bateson, W. (1894). Materials for the Study of Variation Treated With Especial Regard to Discontinuity in the Origin of Species. London, Macmillan. .

de Vries, H. (1901-1903). Die Mutationstheorie. Versuche und Beobachtungen über die Entstehung von Arten im Pflanzenreich. Leipzig, Veit & Co.

Dietrich, M. R. (2003). "Richard Goldschmidt: hopeful monsters and other 'heresies'." Nat Rev Genet 4(1): 68-74.

Eldredge, N. and S. J. Gould (1972). Punctuated equilibria: an alternative to phyletic gradualism. in Models in Paleobiology. T. J. M. Schopf. San Francisco, Freeman, Cooper and Company: 82-115.

Gillham, N. W. (2001). "Evolution by jumps: Francis Galton and William Bateson and the mechanism of evolutionary change." Genetics 159(4): 1383-1392.

Goldschmidt, R. (1940). The Material Basis of Evolution, Reissued (The Silliman Memorial Lectures Series), 1982. New Haven CT, Yale Univ.Press. .

Gould, S. J. (1977). "The Return of Hopeful Monsters." Natural History 86: 22-30. .

Gould, S. J. (1983). "Punctuated Equilibrium and the Fossil Record." Science 219(4584): 439-440.

Gould, S. J. and N. Eldredge (1993). "Punctuated equilibrium comes of age." Nature 366(6452): 223-227.

Gould, S. J. s. (1980). G. G. Simpson, Paleontology and the Modern Synthesis. The Evolutionary Synthesis. E. Mayr and W. B. Provine. Cambridge MA, Harvard University Press: 153–172. .

Kozo-Polyansky, B. M. (1924 (2010, English translation by Victor Fet)). Symbiogenesis: A New Principle of Evolution. Cambridge, Harvard University Press. .

Margulis, L. (1970). Origin of Eukaryotic Cells, Yale Univ. Press. .

Margulis, L. (1971). "Symbiosis and evolution." Sci Am 225(2): 48-57.

Margulis, L. (1981). Symbiosis in Cell Evolution. London, W.H. Freeman Co. .

Margulis, L. and D. Sagan (2002). Acquiring Genomes: A Theory of the Origins of Species. Amherst, MA, Perseus Books Group. .

Mereschkowsky, K. (1910). "Theorie der zwei Plasmaarten als Grundlage der Symbiogenesis, einer neuen Lehre von der Entstehung der Organismen." Biologisches Centralblatt 30: 353367. .

Mereschkowsky, K. (1926). Symbiogenesis and the Origin of Species.

Peterson, E. L. (2008). "William Bateson from Balanoglossus to Materials for the study of variation: the transatlantic roots of discontinuity and the (un)naturalness of selection." J Hist Biol 41(2): 267-305.

Sagan, L. (1967). "On the origin of mitosing cells." J Theor Biol 14(3): 255-274.

Sapp, J. (1994). Evolution by Association: A History of Symbiosis. Oxford, Oxford University Press. .

Sapp, J. (2010). "Saltational symbiosis." Theory Biosci 129(2-3): 125-133.

Simpson G.G (1944). Tempo and Mode in Evolution. New York, Columbia Univ. Press. .

Stebbins, G. L. (1940). "The significance of polyploidy in plant evolution." Am Nat 74: 54–66. .

Stebbins, G. L. (1971). Processes of Organic Evolution. Englewood Cliffs, New Jersey, Prentice-Hall. .

Stebbins, G. L. and F. J. Ayala (1981). "Is a new evolutionary synthesis necessary?" Science 213(4511): 967-971.

Stebbins, J., G.L. (1951). "Cataclysmic Evolution." Scientific American 184(4): 54 –59. .

Wallin, I. E. (1922). "On the nature of mitochondria. I. Observations on mitochondria staining methods applied to bacteria. II. Reactions of bacteria to chemical treatment." American Journal of Anatomy 30(2): 203–229. .

Wallin, I. E. (1922). "On the nature of mitochondria. III. The demonstration of mitochondria by bacteriological methods. IV. A comparative study of the morphogenesis of root-nodule bacteria and chloroplasts." American Journal of Anatomy 30(4): 451–471. .

Wallin, I. E. (1927). Symbionticism and the origin of species. Baltimore, Williams & Wilkins. .