involved, but infrequent chromosome crossovers accounted for the sudden appearance of traits which had been present in the genes all along. Fisher's explanation of polymorphism, 1927 In 1927, Fisher explicitly attacked Punnett's 1915 theory of discontinuous evolution of mimicry. Fisher argued that selection acting on genes making small modifications to the butterfly's phenotype (its appearance) would allow the multiple forms of a polymorphism to be established. Later mutationist theories The understanding that Mendelian genetics could both preserve discrete variations indefinitely, and support continuous variation for natural selection to work on gradually, meant that most biologists from around 1918 onwards accepted natural selection as the driving force of evolution. Mutationism and other alternatives to evolution by natural selection did not however vanish entirely. Berg's nomogenesis, 1922 Lev Berg proposed a combination of mutationism and directed (orthogenetic) evolution in his 1922 book Nomogenesis; or, Evolution Determined by Law. He used evidence from paleontology, zoology, and botany to argue that natural selection had limitations which set a direction for evolution. He claimed that speciation was caused by "mass transformation of a great number of individuals" by directed mass mutations. Willis's macromutations, 1923 In 1923, the botanist John Christopher Willis proposed that species were formed by large mutations, not gradual evolution by natural selection, and that evolution was driven by orthogenesis, which he called "differentiation", rather than by natural selection. Goldschmidt's hopeful monsters, 1940 In his 1940 book The Material Basis of Evolution, the German geneticist Richard Goldschmidt argued for single-step speciation by macromutation, describing the organisms thus produced as "hopeful monsters". Goldschmidt's thesis was universally rejected and widely ridiculed by biologists, who favoured the neo-Darwinian explanations of Fisher, J. B. S. Haldane and Sewall Wright. However, interest in Goldschmidt's ideas has reawakened in the field of evolutionary developmental biology. Nei's mutation-driven evolution, 1987 Contemporary biologists accept that mutation and selection both play roles in evolution; the mainstream view is that while mutation supplies material for selection in the form of variation, all non-random outcomes are caused by natural selection. Masatoshi Nei argues instead that the production of more efficient genotypes by mutation is fundamental for evolution, and that evolution is often mutation-limited. Nei's book received thoughtful reviews; while Wright rejected Nei's thinking as mistaken, Brookfield, Galtier, Weiss, Stoltzfus, and Wagner, although not necessarily agreeing with Nei's position, treated it as a relevant alternative view. Contemporary approaches Reviewing the history of macroevolutionary theories, the