The global phylogeny of the subfamily Sycoryctinae (Pteromalidae): parasites of an obligate mutualism.

The inflorescences of fig trees (Ficus, Moraceae) host well-defined, host plant specific wasp communities that lend themselves to tests of hypotheses on insect diversification. We provide the first estimate of the global molecular phylogeny for the Sycoryctinae - a large subfamily of fig wasps consisting mainly of parasitoids of fig-pollinating wasps. We find strong support for a large Old World clade that contains eight of the eleven genera, in the tribes Sycoryctini and Philotrypesini. The sister taxon is tribe Apocryptini, comprising the genera Apocrypta and Bouceka. Finally, a new tribe, Critogastrini, is raised for the genus Critogaster, sister to all other sycoryctines. At the genus level, we found a general pattern of strong host conservatism, in which closely related wasps associate with closely related figs. Despite this, there is also evidence for multiple host shifts between more distantly related figs in some wasp genera (especially Philotrypesis). We estimate Sycoryctinae to have originated 49-64 Ma, after the initial co-radiation of the host figs and pollinators. Further, conservative assumptions in our analyses probably overestimate the age of the sycoryctines. Together, patterns of host association, evidence for a mix of host constraints and host shifting, and molecular dating suggest that sycoryctine parasites radiated through delayed phylogenetic tracking of their hosts. This contributes to the growing body of literature suggesting that coevolving parasites often radiate after their hosts.