This paper presents the first complete estimate of the phylogenetic relationships among all 197 species of extant and recently extinct ruminants combining morphological, ethological and molecular information. The composite tree is derived by applying matrix representation using parsimony analysis to 164 previous partial estimates, and is remarkably well resolved, containing 159 nodes (>80% of the potential nodes in the completely resolved phylogeny). Bremer decay index has been used to indicate the degree of certainty associated with each clade. The ages of over 80 % of the clades in the tree have been estimated from information in the literature. The supertree for Ruminantia illustrates which areas of ruminant phylogeny are still only roughly known because of taxa with controversial relationships (e.g. Odocoileini, Antilopinae) or not studied in great detail (e.g. Muntiacus). It supports the monophyly of the ruminant families and Pecora. According to this analysis Antilocapridae and Giraffidae constitute the superfamily Giraffoidea, which is the sister group of a clade clustering Bovoidea and Cervoidea. The position of several taxa whose systematic positions have remained controversial in the past (Saiga, Pelea, Aepycerus, Pantholops, Ammotragus, Pseudois) is unambiguously established. Nevertheless, the position of Neotragus and Oreotragus within the original radiation of the non‐bovine bovids remains unresolved in the present analysis. It also shows that six successive rapid cladogenesis events occurred within the infraorder Pecora during the Oligocene to middle Pliocene, which coincided with periods of global climatic change. Finally, the presented supertree will be a useful framework for comparative and evolutionary biologists interested in studies involving the ruminants.

What drives divergence?: Horse evolution has long been held as a classic example of adaptive radiation. It has been thought that an increase in the height of cheek teeth opened up new grass resources, leading to divergence. Cantalapiedra et al. , however, found that although the Equinae have experienced high levels of divergence, these splits do not appear to have been related initially to specific phenotypic changes. Instead, it seems that external environmental drivers and patterns of migration and isolation initiated population divergence, with phenotypic changes emerging once lineages had begun to divide. Science , this issue p. 627

Climatic and environmental shifts have had profound impacts on faunal and floral assemblages globally since the end of the Miocene. We explore the regional expression of these fluctuations in southwestern Europe by constructing long-term records (from ∼11.1 to 0.8 Ma, late Miocene–middle Pleistocene) of carbon and oxygen isotope variations in tooth enamel of different large herbivorous mammals from Spain. Isotopic differences among taxa illuminate differences in ecological niches. The δ¹³C values (relative to VPDB, mean −10.3±1.1‰; range −13.0 to −7.4‰) are consistent with consumption of C₃ vegetation; C₄ plants did not contribute significantly to the diets of the selected taxa. When averaged by time interval to examine secular trends, δ¹³C values increase at ∼9.5 Ma (MN9–MN10), probably related to the Middle Vallesian Crisis when there was a replacement of vegetation adapted to more humid conditions by vegetation adapted to drier and more seasonal conditions, and resulting in the disappearance of forested mammalian fauna. The mean δ¹³C value drops significantly at ∼4.2−3.7 Ma (MN14–MN15) during the Pliocene Warm Period, which brought more humid conditions to Europe, and returns to higher δ¹³C values from ∼2.6 Ma onwards (MN16), most likely reflecting more arid conditions as a consequence of the onset of the Northern Hemisphere glaciation. The most notable feature in oxygen isotope records (and mean annual temperature reconstructed from these records) is a gradual drop between MN13 and the middle Pleistocene (∼6.3−0.8 Ma) most likely due to cooling associated with Northern Hemisphere glaciation.