Centre for Philosophy of Science Applied Evolutionary Epistemology Lab

Folmer Bokma


Course description

The lectures will concern the question to what extent the theory of evolution can be regarded a scientific theory, with particular focus on the question whether the theory of evolution leads to testable predictions. With these questions in mind, we will trace the development of some important aspects of the theory of evolution from before Darwin's time, through Darwin's time and the subsequent rediscovery of Mendelian genetics which sparked the development of the Modern Synthesis, to more recent claims that the Modern Synthesis is in need of revision. The theory of evolution didn't develop in isolation, and the lectures will draw together progress in paleontology, ecology, and especially (quantitative) genetics to provide a solid background for understanding how evolutionary theory developed. Where the lectures touch on philosophical issues there will be scope for group discussion.


Day-by-Day Program

Lecture 1: A Case Study: Deep Flowers for Long Tongues?

  1. Wallace, A. R. 1867. Creation by law. Quarterly Journal of Science 4:471-488.
  2. Deep Flowers for Long Tongues (short discussion in TREE by various authors).
  3. Whittall, J. B. and S. A. Hodges. 2007. Pollinator shifts drive increasingly long nectar spurs in columbine flowers. Nature 447:706-712.
  4. Gould, S. J. and R. C. Lewontin. 1979. The Spandrels of San Marco and the Panglossian Paradigm: A Critique of the Adaptationist Programme. Proceedings of the Royal Society B: Biological Sciences 205:581-598.
  5. (Optional: Nielsen, R. 2009. Adaptationism – 30 years after Gould and Lewontin. Evolution 63-10: 2487–2490).
  6. Bond, S. 2012. Angraecum sesquipedale: Darwin's Great 'Gamble'. M. Brinkworth and F. Weinert (eds.), Evolution 2.0, The Frontiers Collection, Springer-Verlag Berlin Heidelberg.


Lecture 2: Do Species Adapt?

  1. Letter by Hugh Falconer to Darwin; and Darwin's reply (1862).
  2. Bradshaw, A. D. 1991. Genostasis and the limits to evolution. Philosophical Transactions of the Royal Society B: Biological Sciences 333:289-305. (The Croonian Lecture)
  3. Merilä, J. 2012. Evolution in response to climate change: In pursuit of the missing evidence . Bioessays 34: 811–818.


Lecture 3: Constraints on Evolution

  1. (Optional: Haldane, J. B. S. 1957. The Cost of Natural Selection. Journal of Genetics 55:551-524).
  2. Van Valen, L. 1963. Haldane's Dilemma, evolutionary rates, and heterosis. The American Naturalist 47:185-190.
  3. Kimura, M. 1995. Limitations of Darwinian selection in a finite population. Proceedings of the National Academy of Sciences USA 92:2343-2344.
  4. Ohno, S. 1970. Preface to "Evolution by gene duplication". (1/2 page)
  5. Pigliucci, M. 2006. Genetic variance–covariance matrices: a critique of the evolutionary quantitative genetics research program . Biology and Philosophy 21:1–23.
  6. (Optional: Futuyma, D. J. 2010. Evolutionary constraint and ecological consequences. Evolution 64: 1865-1884).


Lecture 4: Speciation: Internal versus External Factors

  1. Cuvier, 1798, article on his "Principe de la correlation des formes".
  2. Riedl, R. 1977. A systems-analytical approach to macro-evolutionary phenomena. Quarterly Review of Biology 52: 351-370.
  3. Eldredge, N. and S. J. Gould. 1972. Punctuated Equilibrium: An alternative to phyletic gradualism. Pages 82-115 in T. J. M. Schopf, ed. Models in Paleobiology. Freeman, Cooper & Co, San Francisco.
  4. Minelli A. and Fusco G. 2012. On the evolutionary developmental biology of speciation. Evolutionary Biology 39: 242-254.
  5. McCarthy, E. M. 2008. On the Origins of New Forms of Life: A New Theory. Chapter 2: On Hybrids.
  6. (Optional: Patterson, N., D. J. Richter, S. Gnerre, E. S. Lander, and D. Reich. 2006. Genetic evidence for complex speciation of humans and chimpanzees. Nature 441:1103-1108).


Lecture 5: Tools for Phylogenetic Macroevolutionary Inferences

  1. Cheverud, J.M., Dow, M.M., and Leutenegger, W. 1985. The quantitative assessment of phylogenetic constraints in comparative analyses: sexual dimorphism in body weight among primates. Evolution 39: 1335-1351.
  2. Mattila, T.M. and Bokma, F. 2008. Extant mammal body masses suggest Punctuated Equilibrium. Proc. Roy. Soc. B. 275:2195-2199.
  3. I will compose further reading material for this lecture (brownian motion; Ornstein-Uhlenbeck; Chapman-Kolmogorov on molecular evolution; branching processes, MCMC, phylogeny construction. Entropy.)