Department of Statistics, University of Oxford, Oxford OX1 3TG, UK and Molecular and Computational Biology Program, Department of Biological Sciences, University of Southern California, CA 90089-2910, USA.|Department of Statistics, University of Oxford, Oxford OX1 3TG, UK and Molecular and Computational Biology Program, Department of Biological Sciences, University of Southern California, CA 90089-2910, USA.|Department of Statistics, University of Oxford, Oxford OX1 3TG, UK and Molecular and Computational Biology Program, Department of Biological Sciences, University of Southern California, CA 90089-2910, USA.|Department of Statistics, University of Oxford, Oxford OX1 3TG, UK and Molecular and Computational Biology Program, Department of Biological Sciences, University of Southern California, CA 90089-2910, USA.|Department of Statistics, University of Oxford, Oxford OX1 3TG, UK and Molecular and Computational Biology Program, Department of Biological Sciences, University of Southern California, CA 90089-2910, USA.