1. A New Look at the Major Features of Evolution Stevan J. Arnold Department of Integrative Biology Oregon State University

2. Outline Criteria for the conservation of individual species G. L. Simpson’s modes of evolution Evolution of body size: data Evolution of body size: models The concept of quantum evolution The concept illustrated with African birds A new criterion for the conservation

3. Criteria for conservation Threatened status Ecological importance (e.g., keystone species) Useful to humans Charisma (e.g., flagship species) Uniqueness (e.g., aardvark)

4. Simpson’s two evolutionary modes: phyletic evolution and quantum evolution Simpson 1944

5. Simpson’s concept of quantum evolution Simpson 1944

6. Modern version of quantum evolution Lande 1976

7. Uyeda et al. 2011

10. ±65% change in body size Uyeda et al. 2011

11. “The Blunderbuss Pattern” Uyeda et al. 2011

12. The multiple-burst model: a process that produces quantum evolution (peak movement, evolution of the lineage mean) Uyeda et al. 2011 A single lineage Time (generations) Lineage mean

13. Burst timing distribution (mean time between bursts = 25 my) White noise distribution (dashed) Burst size distribution (solid) Probability Divergence B C A Interval (years) The model that best fits the data has two modes: phyletic & quantum Uyeda et al. 2011

14. Can we detect quantum evolution in the birds of Central Africa?

15. Provisos Phylogeny ignored Exclusive focus on body size Measure of body size includes tail Focus on Central Africa ignores relatives elsewhere

16. No quantum evolution

17. Quantum evolution of Great Blue Turaco

18. Quantum evolution of ground hornbills

19. Quantum evolution of the Giant Kingfisher

20. Criteria for conservation Threatened status Ecological importance (e.g., keystone species) Useful to humans Charisma (e.g., flagship species) Uniqueness (e.g., aardvark) Replacement time (e.g., ≧ 25 million years for a ground hornbill)

21. Conclusions To account for evolutionary pattern in a large data set we need a model with two modes: phyletic and quantum. Quantum evolutionary events can lead to rapid, substantial evolution but they are rare, with an average waiting time of 25 million years. Some families of birds in Central Africa appear to have experienced quantum evolution (e.g., hornbills, kingfishers, and turacos).

22. References Simpson, G. L. 1944. Tempo and Mode in Evolution. Columbia Univ. Press. Simpson, G. L. 1954. The Major Features of Evolution. Columbia Univ. Press. Lande, R. 1976. Uyeda, J., T. Hansen, S. J. Arnold, J. Pienaar. 2011. The million-year wait for evolutionary bursts. Proceedings of the National Academy of Science. U.S.A. Arnold, S. J. 2014. Phenotypic evolution, the ongoing synthesis. American Naturlist Van Perlo, Ber. 2002. Birds of Western and Central Africa. Princeton Univ. Press.

23. Acknowledgements PhD advisor: Arnold Kluge Research collaborators: Suzanne Estes, Josef Uyeda, Thomas Hansen, Jason Pienaar Data: Phil Gingerich, Andrew Hendry, Michael Kinnison NSF OPUS program: Mark Courtney