Phylogenies are usually dated by calibrating interior nodes against the fossil record. This relies on indirect methods that, in the worst case, misrepresent the fossil information. Here, we contrast such node dating with an approach that includes fossils along with the extant taxa in a Bayesian total-evidence analysis. As a test case, we focus on the early radiation of the Hymenoptera, mostly documented by poorly preserved impression fossils that are difficult to place phylogenetically. Specifically, we compare node dating using nine calibration points derived from the fossil record with total-evidence dating based on 343 morphological characters scored for 45 fossil (4--20 complete) and 68 extant taxa. In both cases we use molecular data from seven markers (∼5 kb) for the extant taxa. Because it is difficult to model speciation, extinction, sampling, and fossil preservation realistically, we develop a simple uniform prior for clock trees with fossils, and we use relaxed clock models to accommodate rate variation across the tree. Despite considerable uncertainty in the placement of most fossils, we find that they contribute significantly to the estimation of divergence times in the total-evidence analysis. In particular, the posterior distributions on divergence times are less sensitive to prior assumptions and tend to be more precise than in node dating. The total-evidence analysis also shows that four of the seven Hymenoptera calibration points used in node dating are likely to be based on erroneous or doubtful assumptions about the fossil placement. With respect to the early radiation of Hymenoptera, our results suggest that the crown group dates back to the Carboniferous, ∼309 Ma (95% interval: 291--347 Ma), and diversified into major extant lineages much earlier than previously thought, well before the Triassic. [Bayesian inference; fossil dating; morphological evolution; relaxed clock; statistical phylogenetics.]

The types and undescribed material of the hymenopteran fossils of the Insect Bed of the Bembridge Marls from the Isle of Wight (UK) are critically revised and studied. A total of 1460 fossils are recorded and attributed to 20 families: Gasteruptiidae s.l. (1); Proctotrupidae (3); Diapriidae (24); Cynipidae (7); Figitidae (6); Pteromalidae (1); Agaonidae (3); Scelionidae (12); Platygastridae (2); Ichneumonidae (32); Braconidae (75); Bethylidae (3); Crabronidae (2); Sphecidae (1); Apidae (2); Scoliidae (1); Tiphiidae (2); Vespidae (4); and Formicidae (1220). Described as new are 51 species, 13 genera, two tribes and two subfamilies. Minimum number of species recorded (either as described species or representing higher taxa with no described species in the assemblage) is 118. The composition of the hymenopteran assemblage is most similar to that of Baltic amber and indicative of a well forested territory, as well as of a humid, equable (aseasonal but not very hot) climate, more typically equable than in the Baltic amber source area, judging from the absence of Aphidiinae and scarcity of aphids.

Long-Lost Pollinators: The rise of angiosperms in the Early Cretaceous (∼140 million years ago) was accompanied by coevolution of a variety of insects, including flies, bees, and wasps required for pollination. Ren et al. (p. 840 ; see the Perspective by Ollerton and Coulthard ) show that three families of scorpionflies had already evolved specialized mouth parts for feeding on the nectar of gymnosperms, as early as the Middle Jurassic (∼170 million years ago). The diversity and specialization of these insects and related plant structures suggests that they were also involved in pollination. These families died out later in the Cretaceous as angiosperms began to dominate.

A total of 5754 ant inclusions from 13 European collections of Baltic, Bitterfeld, Rovno, and Scandinavian ambers are studied and identified as belonging to 147 species, 57 genera, and 9 subfamilies. The taxonomic composition and relative species abundances of species in representative collections are analyzed and considerable differences between the above four types of amber are shown. These differences appear to reflect differences in ecological conditions rather than in age. The Baltic and Bitterfeld ant assemblages are shown to be most similar, the Scandinavian assemblage turns out to be most dissimilar to these, and the Rovno assemblage is shown to be intermediate.