The high extant species diversity of tropical lineages of organisms is usually portrayed as a relatively recent and rapid development or as a consequence of the gradual accumulation or preservation of species over time. These explanations have led to alternative views of tropical forests as evolutionary ''cradles'' or ''museums'' of diversity, depending on the organisms under study. However, biogeographic and fossil evidence implies that the evolutionary histories of diversification among tropical organisms may be expected to exhibit characteristics of both cradle and museum models. This possibility has not been explored in detail for any group of terrestrial tropical organisms. Cephaloleia ͉ diversification ͉ evolutionary radiation ͉ phylogeny T he extraordinarily high species diversity of tropical forest floras and faunas is often attributed to the recent and rapid accumulation of species via high speciation rates (1-4) or the gradual accumulation and͞or preservation of species over time via low extinction rates (5-8). These observations have led to the widespread belief that tropical forests are evolutionary ''cradles'' of diversity for some lineages and ''museums'' of diversity for others (6, 7).Evidence in support of cradle models comes largely from geographic patterns of distribution, species richness, and endemism of extant tropical organisms. These patterns are often ascribed to evolutionary radiation in response to relatively recent climatic, tectonic, or biotic events; e.g., Pleistocene [Ϸ1.8-0.01 mega-annum (Ma) ago] glaciation or Pliocene (Ϸ5.3-1.8 Ma ago) bridging of the Isthmus of Panama (1-4). However, paleontological evidence implies that many of the evolutionary radiations that account for the present diversity (''crown diversification'') of taxonomically disparate groups of tropical organisms may have occurred comparatively early, during the late Paleocene and early to middle Eocene (Thanetian, Ypresian, and Lutetian Ages, Ϸ58.7-40.4 Ma ago) (9-17), associated with global warming, Cenozoic maximum global temperatures (18), and the latitudinal expansion (16, 19) and taxonomic diversification (9,11,14,16,17) of characteristically tropical lineages of plants, consistent with museum models of diversification. Although cradle and museum models are often presented as temporal alternatives (4, 7), their predictions are not mutually exclusive, and, as the aforementioned observations suggest, the evolutionary histories of tropical lineages of organisms may be expected to exhibit features of both kinds of models. However, this possibility has not been explored in detail for any group of terrestrial tropical organisms.Using molecular genetic, paleontologic, and biogeographic evidence, we investigated timing and tempo in the diversification of Cephaloleia, a species-rich genus of herbivorous Neotropical leaf beetles (Chrysomelidae: Cassidinae). Our goals were to determine (i) whether the evolutionary history of Cephaloleia diversification (speciation-extinction) departs significantly from a constant rate m...