Tree balance, time slices, and evolutionary turnover in cretaceous planktonic foraminifera

Studies of phylogenetic tree shape often concentrate on the balance of phylogenies of extant taxa. Paleontological phylogenies (which include extinct taxa) can contain additional useful information and can directly document changes in tree shape through evolutionary time. Unfortunately, the inclusion of extinct taxa lowers the power of direct examinations of tree balance because it increases the range of tree shapes expected under null models of evolution (with equal rates of speciation and extinction across lineages). A promising approach for the analysis of tree shape in paleontological phylogenies is to break the phylogeny down into time slices, examining the shape of the phylogeny of taxa alive at each time slice and changes in that shape between successive time slices. This method was illustrated with 57 time slices through a stratophenetic phylogeny of the Cretaceous planktonic foraminiferal superfamily Globotruncanacea. At 3 of 56 intervals between time slices, 93-92.5 million years ago (MYA), 89-88.5 MYA, and 85.5-84 MYA, the group showed steep increases in imbalance. Although none of these increases were significant after Bonferroni correction, these points in the history of the Globotruncanacea were nevertheless identified as deserving of further macroevolutionary investigation. The 84 MYA time slice coincides with a peak in species turnover for the superfamily. Time slices through phylogenies may prove useful for identifying periods of time when evolution was proceeding in a nonstochastic manner.