Multivariate analyses of skull morphology inform the taxonomy and evolution of geomyoid rodents.

Morphological analyses are critical to quantify phenotypic variation, identify taxa, inform phylogenetic relationships, and shed light on evolutionary patterns. This work is particularly important in groups that display great morphological disparity. Such is the case in geomyoid rodents, a group that includes 2 of the most species-rich families of rodents in North America: the Geomyidae (pocket gophers) and the Heteromyidae (kangaroo rats, pocket mice, and their relatives). We assessed variation in skull morphology (including both shape and size) among geomyoids to test the hypothesis that there are statistically significant differences in skull measurements at the family, genus, and species levels. Our sample includes 886 specimens representing all geomyoid genera and 39 species. We used the geometric mean to compare size across taxa. We used 14 measurements of the cranium and lower jaw normalized for size to compare shape among and within taxa. Our results show that skull measurements enable the distinction of geomyoids at the family, genus, and species levels. There is a larger amount of size variation within Geomyidae than within Heteromyidae. Our phylomorphospace analysis shows that the skull shape of the common ancestor of all geomyoids was more similar to the common ancestor of heteromyids than that of geomyids. Geomyid skulls display negative allometry whereas heteromyid skulls display positive allometry. Within heteromyids, dipodomyines, and non-dipodomyines show significantly different allometric patterns. Future analyses including fossils will be necessary to test our evolutionary hypotheses.

Craniodental divergence associated with bite force between hybridizing pine squirrels (Tamiasciurus).

Bite force can be a limiting factor in foraging and can significantly affect the competitive ability and lifetime fitness of mammals. Tamiasciurus squirrels feed primarily on conifer seeds and have a strong bite force to mechanically extract seeds from conifer cones with their mouths. In the North Cascades region, Douglas squirrels (Tamiasciurus douglasii) and red squirrels (T. hudsonicus) occupy ecologically different forests with different hardnesses in conifer cones. The ranges of these species overlap in a narrow hybrid zone where these forests meet near the crest of the North Cascades. We examined interspecific divergence in dietary ecomorphology in allopatry, in sympatry within the hybrid zone, and between hybrids and each parental species. We focused on three craniodental traits, including the incisor-strength index as a proxy measure for maximal bite force, cranial-suture complexity, and mandible shape. We find that these sister squirrel species differ in bite force and suture complexity in allopatry and sympatry and that mandible shape changes with the expected hardness of accessed food items, but is not significantly different between species. Furthermore, we find that hybrids display morphologies that overlap with hybrid zone red squirrels but not with hybrid zone Douglas squirrels. This work shows how important ecological processes at shallow evolutionary timescales can impact the divergence of morphological traits in taxa with extreme conservation of craniomandibular shape.

The earliest dipodomyine heteromyid in North America and the phylogenetic relationships of geomorph rodents.

Dipodomyine heteromyids (kangaroo rats and mice) are a diverse group of arid-adapted ricochetal rodents of North America. Here, a new genus and species of a large dipodomyine is reported from early Miocene-aged deposits of the John Day Formation in Oregon that represents the earliest record of the subfamily. The taxon is known from a single specimen consisting of a nearly complete skull, dentary, partial pes, and caudal vertebra. The specimen is characterized by a mosaic of ancestral and highly derived cranial features of heteromyids. Specifically, the dental morphology and some cranial characteristics are similar to early heteromyids, but other aspects of morphology, including the exceptionally inflated auditory bullae, are more similar to known dipodomyines. This specimen was included in a phylogenetic analysis comprising 96 characters and the broadest sampling of living and extinct geomorph rodents of any morphological phylogenetic analysis to date. Results support the monophyly of crown-group Heteromyidae exclusive of Geomyidae and place the new taxon within Dipodomyinae. The new heteromyid is the largest known member of the family. Analyses suggest that large body size evolved several times within Heteromyidae. Overall, the morphology of the new heteromyid supports a mosaic evolution of the open-habitat adaptations that characterize kangaroo rats and mice, with the inflation of the auditory bulla appearing early in the group, and bipedality/ricochetal locomotion appearing later. We hypothesize that cooling and drying conditions in the late Oligocene and early Miocene favored adaptations for life in more open habitats, resulting in increased locomotor specialization in this lineage over time from a terrestrial ancestor.

Sexual dimorphism in cranial shape and size in geomyoid rodents: multivariate and evolutionary perspectives.

Geomyoid rodents provide a great study system for the analysis of sexual dimorphism. They are polygynic and many inhabit harsh arid environments thought to promote sexual dimorphism. In fact, there has been extensive work published on the sexual size dimorphism of individual populations and species within this rodent clade. However, little work has been undertaken to assess the evolutionary patterns and processes associated with this sexual dimorphism. We use multivariate analyses of cranial measurements in a phylogenetic framework to determine the distribution of size and shape dimorphism among geomyoids and test for Rensch's rule. Our results suggest that sexual dimorphism is more common in geomyids than heteromyids, but it is not in fact universal. There is evidence for variation in sexual dimorphism across populations. Additionally, in many taxa, geographic variation appears to overwhelm existing sexual dimorphism. We find support for the repeated independent evolution of shape and size dimorphism across geomyoid taxa, but we do not find support for an association between size and shape dimorphism. There is no evidence for Rensch's rule in geomyoids, whether at the superfamily or family level. Together, our findings suggest that there is no single explanation for the evolution of sexual dimorphism in geomyoids and that, instead, it is the product of numerous evolutionary events. Future studies incorporating phylogenetic relationships will be necessary to paint a more complete picture of the evolution of sexual dimorphism in geomyoids.

The oldest semi-aquatic beaver in the world and a new hypothesis for the evolution of locomotion in Castoridae.

The North American rodent fossil record includes hundreds of species representing both an incredible taxonomic diversity and great ecological disparity. Although it is during the Oligocene that taxonomic diversity first peaks, it is not until the Miocene, almost 10 Myr later, that many ecologies, particularly locomotory ecologies, are recorded. Here, I present a new Oligocene-aged species of beaver from Montana, Microtheriomys articulaquaticus sp. nov., which represents the oldest semi-aquatic rodent in North America and the oldest amphibious beaver in the world, pushing the advent of semi-aquatic ecology in beavers by 7 Myr. I also provide morphological data supporting a terrestrial ecology for the sister taxon to Castoridae. Together with existing data, these findings lead to a new hypothesis for the evolutionary ecology of castorids whereby swimming was exapted from burrowing during the Oligocene. This evolution of semi-aquatic locomotion may have taken place in North America instead of Eurasia. It started in small beavers with gigantism achieved only much later. Indeed, body size evolution in castoroids follows a directional drift. Beavers obey Cope's rule, a selection for larger size over time that appears associated with semi-aquatic ecology and may well explain their low modern diversity.

Ecological correlates of the morphology of the auditory bulla in rodents: Application to the fossil record.

For rodents, hearing is essential to survival. It enables predator evasion, prey detection, and conspecific recognition; it is also likely to be constrained by the physical environment. The resulting hypothetical link between tympanic bulla morphology and ecology has never been investigated across a broad array of rodent species before. Such link may enable the determination of the ecological affinities of many fossil species only known from partial skulls. In this study, we used geometric morphometrics to quantify the shape of the auditory bulla of 197 specimens representing 91 species from 17 families of extant rodents across four different locomotory modes. We used landmarks and semi-landmarks on the ventral and lateral views of the skull to capture morphological characteristics of the bulla and external auditory meatus (EAM). Our results demonstrate an association between bullar morphology and locomotion in rodents. Bullar shape enables the correct classification of 76% of the species in our training set. Fossorial taxa, in particular, show a characteristic morphology including an asymmetric bulla with a dorsally located and laterally expanded EAM that has a small opening diameter. A phylogenetically informed flexible discriminant analysis shows a weak phylogenetic effect on tympanic morphology. There is no evidence for differences in bullar hypertrophy across locomotory categories. The application of this approach to select fossil rodents from the Oligo-Miocene shows broad agreements with prior studies and yields new locomotory inferences for 14 fossil species, including the first proposed locomotion for members of the family Florentiamyidae. Such results call for the timing of burrowing diversification in rodents to be reevaluated.

Locomotory adaptations in entoptychine gophers (Rodentia: Geomyidae) and the mosaic evolution of fossoriality.

Pocket gophers (family Geomyidae) are the dominant burrowing rodents in North America today. Their fossil record is also incredibly rich; in particular, entoptychine gophers, a diverse extinct subfamily of the Geomyidae, are known from countless teeth and jaws from Oligocene and Miocene-aged deposits of the western United States and Mexico. Their postcranial remains, however, are much rarer and little studied. Yet, they offer the opportunity to investigate the locomotion of fossil gophers, shed light on the evolution of fossoriality, and enable ecomorphological comparisons with contemporaneous rodents. We present herein a quantitative study of the cranial and postcranial remains of eight different species of entoptychine gophers as well as many contemporary rodent species. We find a range of burrowing capabilities within Entoptychinae, including semifossorial scratch-digging animals and fossorial taxa with cranial adaptations to burrowing. Our results suggest the repeated evolution of chisel-tooth digging across genera. Comparisons between entoptychine gophers and contemporaneous rodent taxa show little ecomorphological overlap and suggest that the succession of burrowing rodent taxa on the landscape may have had more to do with habitat partitioning than competition.

The first tetrapod from the mid-Miocene Clarkia lagerstätte (Idaho, USA).

The Clarkia lagerstätte (Latah Formation) of Idaho is well known for its beautifully preserved plant fossils as well as a fauna of insects and fish. Here we present the first known tetrapod fossil from these deposits. This specimen, recovered from the lower anoxic zone of the beds, is preserved as a carbonaceous film of a partial skeleton associated with a partial lower incisor and some tooth fragments. The morphology of the teeth indicates that the first tetrapod reported from Clarkia is a rodent. Its skeletal morphology as well as its bunodont and brachydont dentition suggests that it is a member of the squirrel family (Sciuridae). It is a large specimen that cannot be assigned to a known genus. Instead, it appears to represent the first occurrence of a new taxon with particularly gracile postcranial morphology likely indicative of an arboreal ecology. This new specimen is a rare glimpse into the poorly known arboreal mammal fossil record of the Neogene. It supports a greater taxonomic and ecological diversity of Miocene Sciuridae than previously recognized and offers new lines of inquiry in the paleoecological research enabled by the unique preservation conditions of the Clarkia biota.