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Download paper Released 2008-06-28

Evolution beyond the 20th Century paradigm

Biotic evolution unfolds from self-replication, just as the Universe unfolds from Big Bag

Evolution by natural selection is a history that unfolds from self-replication; just as the Universe unfolds from Big Bang. But the theory of evolution by natural selection was originally constructed the other way around; backwards, by letting natural selection be contingent upon evolutionary history. Instead of providing a mathematical theory that explained observed biotic patterns from first principles of self-replication, Darwin proposed that species are understood from their evolutionary history of common origin, their heritable variation, and a struggle among individuals for existence.

The approach of defining natural selection from evolutionary history became the well-established tradition in the 20th Century. In a strong interpretation, Gould (2002) argued that biological organisation cannot in principle be predicted forwards even with full knowledge of antecedent conditions, but can only be understood backwards after time's actual unfolding. Major evolutionary transitions were seen as evolutionary possibilities (Maynard Smith and Szathmary, 1995), and the very notion that biology might harbour ahistorical universal laws other than “chance and necessity” became simple nonsense (Kauffman, 1993).

The theory of Malthusian Relativity was constructed as an alternative to evolutionary contingency; to show that biotic evolution is indeed an inevitable unfolding from the natural selection that follows from the origin of self-replicating entities. This suggests that the strong interpretation of evolutionary contingency reflected a young scientific discipline that had not yet discovered the underlying Laws of regularity.

In “Inevitable evolution: back to The Origin and beyond the 20th Century paradigm of contingent evolution by historical natural selection”, a paper in Biological Reviews, I cover the development of Malthusian Relativity from 1995 to 2008. It describes differences and similarities between the historical and deterministic selection processes. It illustrates concepts using life history models on large body masses and limited reproductive rates, reviews evolution with a wider focus on major evolutionary trends and transitions, and shows that biotic evolution is driven by a universal natural selection.

Given suitable environmental conditions, the paper shows that selection by net energy and density dependent competitive interactions unfolds to higher-level selection for life history transitions from the simple asexually reproducing self-replicator to large bodied organisms with senescence and sexual reproduction between males and females, and in some cases, to the fully evolved eusocial colony with thousands of offspring workers. This defines an evolutionary arrow of time for open thermodynamic systems with a constant inflow of energy, predicting similar routes for long-term evolution on similar planets.


  • Gould, S.J. 2002. The structure of evolutionary theory. The Belknap Press of Harvard University Press, Cambridge.
  • Kauffman, S.A. 1993. The origins of order. Self-organisation and selection in evolution. Oxford University Press, New York.
  • ry, 1995Maynard:Smith:Szathmary:1995MaynardSmith, J., and E.Szathm\'ary 1995. The major transitions in evolution. W.H. Freeman Spektrum, Oxford.