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Four groups of equids, “Anchitheriinae,” Merychippine-grade Equinae, Hipparionini, and Equini, coexisted in the middle Miocene, but only the Equini remains after 16 Myr of evolution and extinction. Each group is distinct in its occlusal enamel pattern. These patterns have been compared qualitatively but rarely quantitatively. The processes influencing the evolution of these occlusal patterns have not been thoroughly investigated with respect to phylogeny, tooth position, and climate through geologic time. We investigated Occlusal Enamel Index, a quantitative method for the analysis of the complexity of occlusal patterns. We used analyses of variance and an analysis of co-variance to test whether equid teeth increase resistive cutting area for food processing during mastication, as expressed in occlusal enamel complexity, in response to increased abrasion in their diet. Results suggest that occlusal enamel complexity was influenced by climate, phylogeny, and tooth position through time. Occlusal enamel complexity in middle Miocene to Modern horses increased as the animals experienced increased tooth abrasion and a cooling climate.

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Enamel patterns on the occlusal surfaces of equid teeth are asserted to have tribal-level differences. The most notable example compares the Equini and Hipparionini, where Equini have higher crowned teeth with less enamel-band complexity and less total occlusal enamel than Hipparionini. Whereas previous work has successfully quantified differences in enamel band shape by dividing the length of enamel band by the square root of the occlusal surface area (Occlusal Enamel Index, OEI), it was clear that OEI only partially removes the effect of body size. Because enamel band length scales allometrically, body size still has an influence on OEI, with larger individuals having relatively longer enamel bands than smaller individuals. Fractal dimensionality (D) can be scaled to any level, so we have used it to quantify occlusal enamel complexity in a way that allows us to get at an accurate representation of the relationship between complexity and body size. To test the hypothesis of tribal-level complexity differences between Equini and Hipparionini, we digitally traced a sample of 98 teeth, one tooth per individual; 31 Hipparionini and 67 Equini. We restricted our sampling to the P3-M2 to reduce the effect of tooth position. After calculating the D of these teeth with the fractal box method which uses the number of boxes of various sizes to calculate the D of a line, we performed a t-test on the individual values of D for each specimen, comparing the means between the two tribes, and a phylogenetically informed generalized least squares regression (PGLS) for each tribe with occlusal surface area as the independent variable and D as the dependent variable. The slopes of both PGLS analyses were compared using a t-test to determine if the same linear relationship existed between the two tribes. The t-test between tribes was significant (p < 0.0001), suggesting different D populations for each lineage. The PGLS for Hipparionini was a positive but not significant (p = 0.4912) relationship between D and occlusal surface area, but the relationship for Equini was significantly negative (p = 0.0177). λ was 0 for both tests, indicating no important phylogenetic signal is present in the relationship between these two characters, thus the PGLS collapses down to a non-phylogenetic generalized least squares (GLS) model. The t-test comparing the slopes of the regressions was not significant, indicating that the two lineages could have the same relationship between D and occlusal surface area. Our results suggest that the two tribes have the same negative relationship between D and occlusal surface area but the Hipparionini are offset to higher values than the Equini. This offset reflects the divergence between the two lineages since their last common ancestor and may have constrained their ability to respond to environmental change over the Neogene, leading to the differential survival of the Equini.

Abstract.-As many as eight species of the "anchitherine" equid Miohippus have been identified from the John Day Formation of Oregon, but no statistical analysis of variation in these horses has yet been conducted to determine if that level of diversity is warranted. Variation of the anterior-posterior length and transverse width of upper and lower teeth of Turtle Cove Member Miohippus was compared to that of M. equinanus, Mesohippus bairdii, Equus quagga, and Tapirus terrestris using t tests of their coefficients of variation (V). None of the t tests are significant, indicating that the variation seen in Turtle Cove Miohippus is not significantly different from any of the populations of other perissodactyls examined in this study. Data also indicate that Mesohippus is present in the Turtle Cove Member. Additionally, hypostyle condition, used to diagnose all species of Miohippus, was found to be related to stage of wear using an ordered logistic regression. Only two species of equid, one Miohippus and one Mesohippus, in the Turtle Cove Member can be identified, therefore only Miohippus annectens, the genotype and first species described from the region, can be recognized as the sole Miohippus species known from the Turtle Cove assemblage. There are insufficient data to determine which species of Mesohippus is present. The dependence of hypostyle condition on crown height in Miohippus implies that wear stage must also be considered in investigations of dental morphology in the "Anchitheriinae."

A recently identified left dentary of Harpagolestes cf. uintensis represents the first mesonychid material known from the Pacific Northwest. The specimen is from the Hancock Quarry (Clarno Unit, John Day Fossil Beds National Monument), which is in the uppermost subunit of the Clarno Formation (middle Eocene, ~40 Ma). The sediments of the Hancock Quarry were deposited by a meandering river system during the middle Eocene when north-central Oregon had a subtropical climate. As with many other mammals from the Hancock Quarry, Harpagolestes participated in an Asian-North American faunal interchange; species of Harpagolestes are known from the Eocene of both continents. Harpagolestes was carnivorous, and members of the genus were likely bone-crushers. Characteristic bone-crushing wear is visible on the occlusal surfaces of the Hancock Quarry specimen's premolars and molars. With the aid of CT scans, it has been determined that the Hancock Quarry Harpagolestes contains the alveoli for c1, p1-2, and m3, and preserves the crowns of p3-4 and m1-2. The molariform teeth have a large, conical trigonid with a bulbous talonid. The protoconid of p3 and p4 is tilted posteriorly. This specimen of Harpagolestes cf. uintensis represents a new large carnivore in the Hancock Quarry ecosystem, adds to the known diversity of the Oregon middle Eocene, and is the only known occurrence of a mesonychid in the Pacific Northwest.

A paleontological deposit near San Clemente de Térapa represents one of the very few Rancholabrean North American Land Mammal Age sites within Sonora, Mexico. During that time, grasslands were common, and the climate included cooler and drier summers and wetter winters than currently experienced in northern Mexico. Here, we demonstrate restructuring in the mammalian community associated with environmental change over the past 40,000 years at Térapa. The fossil community has a similar number of carnivores and herbivores whereas the modern community consists mostly of carnivores. There was also a 97% decrease in mean body size (from 289 kg to 9 kg) because of the loss of megafauna. We further provide an updated review of ungulates and carnivores, recognizing two distinct morphotypes of Equus, including E. scotti and a slighter species; as well as Platygonus compressus; Camelops hesternus; Canis dirus; and Lynx rufus; and the first regional records of Palaeolama mirifica, Procyon lotor, and Smilodon cf. S. fatalis. The Térapa mammals presented here provide a more comprehensive understanding of the faunal community restructuring that occurred in northern Mexico from the late Pleistocene to present day, indicating further potential biodiversity loss with continued warming and drying of the region.