Nicholas A. Famoso

Occlusal Enamel Complexity in Middle Miocene to Holocene Equids (Equidae: Perissodactyla) of North America

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.

Are Hypsodonty and Occlusal Enamel Complexity Evolutionarily Correlated in Ungulates

The spread of grasslands and cooling climate in the Miocene contributed to an increasingly abrasive diet for ungulates. This increase in abrasiveness is proposed to select for both hypsodonty and increasing complexity of occlusal enamel bands. If these traits evolved in response to strong selection to resist tooth wear while feeding in grassland habitats, we might expect them to have evolved in a correlated fashion. If, on the other hand, there was a developmental or physiological constraint, or if selection was not strong on total enamel production, we would expect species to have evolved one or the other of these traits at a time, producing an uncorrelated, or even inversely correlated, pattern of trait evolution. To test these hypotheses, we examined the Occlusal Enamel Index (OEI) and Hypsodonty Index (HI) of 773 ungulate teeth. We tested the dependence of OEI on HI for the orders Artiodactyla and Perissodactyla using phylogenetic generalized least squares regression (PGLS). The two traits are not significantly correlated in the PGLS, for Artiodactyla and Perissodactyla. Despite their physical proximity, close functional utility, and conventional correlation, our results reject the hypothesis that HI and OEI are evolutionarily linked in these lineages, suggesting that selection to resist tooth wear was not so strong as to drive the overall evolutionary trajectory of both these traits at the same time.

On the relationship between enamel band complexity and occlusal surface area in Equids (Mammalia, Perissodactyla)

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.

Statistical analysis of dental variation in the Oligocene equid Miohippus (Mammalia, Perissodactyla) of Oregon

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.”

Correction to the Holotype (AMNH FM 9394) of Merychippus proparvulus Osborn, 1918 (Perissodactyla, Equidae)

On a recent research trip to the American Museum of Natural History (AMNH), it was discovered that the holotype of Merychippus proparvulus Osborn, 1918:117 (Perissodactyla, Equidae), included postc...