vol. 196 no. 2 509-522
- Received October 2, 2013.
- Accepted November 28, 2013.
- Published online February 4, 2014.
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- *Department of Biology, University of Pennsylvania, Philadelphia, Pennsylvania 19104
- †Department of Organismic and Evolutionary Biology and FAS Center for Systems Biology, Harvard University, Cambridge, Massachusetts 02138
- ↵2Corresponding author: Department of Biology, University of Pennsylvania, Philadelphia, PA 19104. E-mail: