The role of fossils in dating the tree of life has been misunderstood. Fossils can provide good "minimum" age estimates for branches in the tree, but "maximum" constraints on those ages are poorer. Current debates about which are the "best" fossil dates for calibration move to consideration of the most appropriate constraints on the ages of tree nodes. Because fossil-based dates are constraints, and because molecular evolution is not perfectly clock-like, analysts should use more rather than fewer dates, but there has to be a balance between many genes and few dates versus many dates and few genes. We provide "hard" minimum and "soft" maximum age constraints for 30 divergences among key genome model organisms; these should contribute to better understanding of the dating of the animal tree of life.

SYNOPSIS Early dinosaur evolution has been the subject of several phylogenetic studies and the position of certain basal forms is currently debated. This is the case for the oldest known members of the group, excavated from the Late Triassic Ischigualastian beds of South America, such as Herrerasaurus, Eoraptor, Pisanosaurus, Saturnalia and Staurikosaurus. A new cladistic analysis of the early dinosaur radiation was performed to assess the relationships among the three major clades (Ornithischia, Sauropodomorpha and Theropoda) and to define the phylogenetic position of the basal members of the group. The most parsimonious hypothesis has Silesaurus opolensis as the sister taxon to a dichotomy including monophyletic Saurischia and Ornithischia. The latter includes Pisanosaurus mertii, and the former all other well-known Triassic dinosaurs. Saurischia is composed of two major monophyletic groups: Herrerasauridae (including Herrerasaurus ischigualastensis and Staurikosaurus pricei) and Eusaurischia (including the theropod and sauropodomorph lineages), while Eoraptor lunensis appears to represent the sister taxon to Eusaurischia. Saturnalia tupiniquim is a stem-taxon to Sauropodomorpha and Guaibasaurus candelariensis might belong to the theropod branch. Some of these hypotheses are, however, not strongly supported. Especially uncertain are the affinities of Silesaurus and Guaibasaurus. The latter can only be safely regarded as a saurischian, while the former might belong to the ornithischian lineage. The dinosaurian affinities of Eoraptor and Herrerasauridae are strongly supported. Yet, the possibility that they (especially Eoraptor) represent basal theropods, rather than basal saurischians, cannot be dismissed. In fact, basal saurischian evolution is still too poorly understood for a definitive hypothesis of relationships to be presented.

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The rise and diversification of the dinosaurs in the Late Triassic, from 230 to 200 million years ago, is a classic example of an evolutionary radiation with supposed competitive replacement. A comparison of evolutionary rates and morphological disparity of basal dinosaurs and their chief "competitors," the crurotarsan archosaurs, shows that dinosaurs exhibited lower disparity and an indistinguishable rate of character evolution. The radiation of Triassic archosaurs as a whole is characterized by declining evolutionary rates and increasing disparity, suggesting a decoupling of character evolution from body plan variety. The results strongly suggest that historical contingency, rather than prolonged competition or general "superiority," was the primary factor in the rise of dinosaurs.

Evolution may be dominated by biotic factors, as in the Red Queen model, or abiotic factors, as in the Court Jester model, or a mixture of both. The two models appear to operate predominantly over different geographic and temporal scales: Competition, predation, and other biotic factors shape ecosystems locally and over short time spans, but extrinsic factors such as climate and oceanographic and tectonic events shape larger-scale patterns regionally and globally, and through thousands and millions of years. Paleobiological studies suggest that species diversity is driven largely by abiotic factors such as climate, landscape, or food supply, and comparative phylogenetic approaches offer new insights into clade dynamics.

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The phylogenetic position of Scleromochlus taylori has been disputed recently, in terms of whether it is a basal sister group of Pterosauria or of Dinosauromorpha. The seven specimens, all from the Lossiemouth Sandstone Formation (late Carnian, Late Triassic) of Lossiemouth, near Elgin, north–east Scotland, suggest that Scleromochlus shares no unique features with either Pterosauria or Dinosauromorpha, together the Ornithodira, but is a close outgroup. Scleromochlus retains a primitive ankle structure, and it has a slender humerus, femur and fibula. Scleromochlus shows the classic bird–like characters of a tibia that is longer than the femur, and a closely appressed group of four elongate metatarsals. A new group name, Avemetatarsalia (‘bird feet’), is established here for the clade consisting of Scleromochlus and Ornithodira, and their descendants. A reanalysis of crown–group archosaur relationships confirms the split into Crurotarsi (crocodile relatives) and Ornithodira (bird relatives), as well as the clear division of Ornithodira into Pterosauria and Dinosauromorpha. Relationships within Crurotarsi are, however, much less clear: Ornithosuchidae probably reside within that clade, and there might be a clade ‘Rauisuchia’ consisting of Prestosuchidae and Postosuchus , but support for these relationships is weak. Scleromochlus was probably a bipedal cursor that could adopt a digitigrade stance. However, it is possible that Scleromochlus was also a saltator, capable of leaping long distances.

Analysis of the fossil record of microbes, algae, fungi, protists, plants, and animals shows that the diversity of both marine and continental life increased exponentially since the end of the Precambrian. This diversification was interrupted by mass extinctions, the largest of which occurred in the Early Cambrian, Late Ordovician, Late Devonian, Late Permian, Early Triassic, Late Triassic, and end-Cretaceous. Most of these extinctions were experienced by both marine and continental organisms. As for the periodicity of mass extinctions, no support was found: Seven mass extinction peaks in the last 250 million years are spaced 20 to 60 million years apart.