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Some molecular clock estimates of divergence times of taxonomic groups undergoing evolutionary radiation are much older than the groups' first observed fossil record. Mathematical models of branching evolution are used to estimate the maximal rate of fossil preservation consistent with a postulated missing history, given the sum of species durations implied by early origins under a range of species origination and extinction rates. The plausibility of postulated divergence times depends on origination, extinction, and preservation rates estimated from the fossil record. For eutherian mammals, this approach suggests that it is unlikely that many modern orders arose much earlier than their oldest fossil records.

The hypocone, a cusp added to the primitively triangular upper molar teeth of therian mammals, has evolved convergently >20 times among mammals during the Cenozoic. Acquisition of the hypocone itself involves little phenotypic change, but subsequent diversification of groups possessing the hypocone may be greatly enhanced. Our analysis of the Cenozoic mammalian radiations, including the Recent fauna, shows that high species diversity of mammals with hypocones and association of the hypocone with herbivory strongly support recognition of the hypocone as a key innovation that has allowed invasion of, and diversification within, herbivorous adaptive zones. In contrast, mammals lacking hypocones show no marked increase in species diversity during the Cenozoic.occlusal contact between metacrista and paracristid (16), crests that are well developed in carnivorous mammals. Thus, possession of a hypocone is generally thought to be associated with herbivory.We investigated the evolutionary potential of the hypocone in two ways. First, we compared taxonomic diversity of living, terrestrial, nonvolant mammals with different molar types across dietary groups to determine whether the hypocone is indeed associated with herbivorous habits and whether mammals with hypocones are more diverse today than those lacking hypocones. Second, we compared the taxonomic diversity of fossil mammals with different molar types across geologic time to determine whether the presence of a hypocone has been associated with higher standing diversity and radiations on each continent.

A classic example of adaptive radiation is the diversification of Cenozoic ungulates into herbivore adaptive zones. Their taxonomic diversification has been associated with changes in molar tooth morphology. Analysis of molar crown types of the Artiodactyla, Perissodactyla, and archaic ungulates ("Condylarthra") shows that the diversity of genera and crown types was high in the Eocene. Post-Eocene molars of intermediate crown types are rare, and thus the ungulate fauna contained more taxa having fewer but more disparate crown types. Taxonomic diversity trends alone give incomplete descriptions of adaptive radiations.

BackgroundThe patterning cascade model of tooth morphogenesis accounts for shape development through the interaction of a small number of genes. In the model, gene expression both directs development and is controlled by the shape of developing teeth. Enamel knots (zones of nonproliferating epithelium) mark the future sites of cusps. In order to form, a new enamel knot must escape the inhibitory fields surrounding other enamel knots before crown components become spatially fixed as morphogenesis ceases. Because cusp location on a fully formed tooth reflects enamel knot placement and tooth size is limited by the cessation of morphogenesis, the model predicts that cusp expression varies with intercusp spacing relative to tooth size. Although previous studies in humans have supported the model's implications, here we directly test the model's predictions for the expression, size, and symmetry of Carabelli cusp, a variation present in many human populations.Methodology/Principal FindingsIn a dental cast sample of upper first molars (M1s) (187 rights, 189 lefts, and 185 antimeric pairs), we measured tooth area and intercusp distances with a Hirox digital microscope. We assessed Carabelli expression quantitatively as an area in a subsample and qualitatively using two typological schemes in the full sample. As predicted, low relative intercusp distance is associated with Carabelli expression in both right and left samples using either qualitative or quantitative measures. Furthermore, asymmetry in Carabelli area is associated with asymmetry in relative intercusp spacing.Conclusions/SignificanceThese findings support the model's predictions for Carabelli cusp expression both across and within individuals. By comparing right-left pairs of the same individual, our data show that small variations in developmental timing or spacing of enamel knots can influence cusp pattern independently of genotype. Our findings suggest that during evolution new cusps may first appear as a result of small changes in the spacing of enamel knots relative to crown size.

Field surveys of the Upper Cretaceous Hell Creek Formation in southwesternNorth Dakota since 1986 have produced a total of 10 124 specimens from 42 vertebrate microsites and an additional 41 dinosaur skulls or partial skeletons or skulls from separate sites representing 61 taxa of vertebrates dominated by fish, dinosaurs, turtles, and crocodilians. Common elements of this diverse fauna occur to within 2.37 m of the Cretaceous-Tertiary (K-T) boundary. The stratigraphically highest fossil in the study is a partial ceratopsian skeleton 1.77 m below the K-T boundary in the basal Fort Union Formation. All dinosaurs that occur at more than two sites also occur at the highest level that yielded more than 500 specimens (8.40 m below the K-T boundary). The fine-grained uppermost 2 m of the Hell Creek is nearly devoid of all fossils, including taxa known to occur in the overlying formation. The absence of channel deposits in this part of the formation may be the reason for the absence of fossil localities. The presence of marine-tolerant taxa suggests that the study area may have been adjacent to a previously unidentified seaway of latest Cretaceous age. Rarefaction analysis indicates no evidence for a decline in vertebrate diversity through the formation or dinosaurian diversity in the 3 m below the K-T boundary. Our results are not compatible with gradual vertebrate extinction at the end of the Cretaceous.

The patterning cascade model of tooth morphogenesis has emerged as a useful tool in explaining how tooth shape develops and how tooth evolution may occur. Enamel knots, specialized areas of dental epithelium where cusps initiate, act as signaling centers that direct the growth of surrounding tissues. For a new cusp to form, an enamel knot must form beyond the inhibition fields of other enamel knots. The model predicts that the number and size of cusps depends on the spacing between enamel knots, reflected in the spacing between cusps. Recently, work by our group demonstrated that the model predicted Carabelli trait expression in human first molars. Here we test whether differences in Carabelli trait expression along the molar row can also be predicted by the model. Crown areas and intercusp distances were measured from dental casts of 316 individuals with a digital microscope. Although absolute cusp spacing is similar in first and second molars, the smaller size and more triangular shape of second molars results in larger cusp spacing relative to size and, likely, less opportunity for the Carabelli trait to form. The presence and size of the hypocone (HY) and a range of small accessory cusps in a larger sample of 340 individuals were also found to covary with the Carabelli trait in a complex way. The results of this study lend further support to the view that the dentition develops, varies, and evolves as a single functional complex.

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