Danielle Fraser

Controlled feeding trials with ungulates: a new application of in vivo dental molding to assess the abrasive factors of microwear

ABSTRACT Microwear, the quantification of microscopic scratches and pits on the occlusal surfaces of tooth enamel, is commonly used as a paleodietary proxy. For ungulates (hoofed mammals), scratch-dominant microwear distinguishes modern grazers from browsers, presumably as a result of abrasion from grass phytoliths (biogenic silica). However, it is also likely that exogenous grit (i.e. soil, dust) is a contributing factor to these scratch-dominant patterns, which may reflect soil ingestion that varies with feeding height and/or environmental conditions (e.g. dust production in open and/or arid habitats). This study assessed the contribution of exogenous grit to tooth wear by measuring the effects of fine- and medium-grained silica sand on tooth enamel using a novel live-animal tooth-molding technique. It therefore constitutes the first controlled feeding experiment using ungulates and the first in vivo experiment using abrasives of different sizes. Four sheep were fed three diet treatments: (1) a mixture of Garrison and Brome hay (control), (2) hay treated with fine-grained silica sand (180–250 µm) and (3) hay treated with medium-grained silica sand (250–425 µm). We found a significant increase in pit features that was correlated with an increase in grain size of grit, corroborating earlier chewing simulation experiments that produced pits through grit-induced abrasion (i.e. the ‘grit effect’). Our results support an interpretation of large silica grains fracturing to create smaller, more abundant angular particles capable of abrasion, with jaw movement defining feature shape (i.e. scratch or pit). Summary: Controlled feeding trials illustrate the effects of exogenous grit on the microscopic tooth wear of hoofed mammals, with implications for paleoecological analyses.

Comparing ungulate dietary proxies using discriminant function analysis

A variety of tooth‐wear and morphological dietary proxies have been proposed for ungulates. In turn, they have been applied to fossil specimens with the purpose of reconstructing the diets of extinct taxa. Although these dietary proxies have been used in isolation and in combination, a consistent set of statistical analyses has never been applied to all of the available datasets. The purpose of this study is to determine how well the most commonly used dietary proxies classify ungulates as browsers, grazers, and mixed feeders individually and in combination. Discriminant function analysis is applied to individual dietary proxies (hypsodonty, mesowear, microwear, and several cranial dietary proxies) and to combinations thereof to compare rates of successful dietary classification. In general, the tooth‐wear dietary proxies (mesowear and microwear) perform better than morphological dietary proxies, though none are strong proxies in isolation. The success rates of the cranial dietary proxies are not increased substantially when ruminants and bovids are analyzed separately, and significance among the three dietary guilds is reduced when controlling for phylogenetic relatedness. The combination of hypsodonty, mesowear, and microwear is found to have a high rate of successful dietary classification, but a combination of all commonly used proxies increases the success rate to 100%. In most cases, mixed feeders bear the greatest resemblance to browsers suggesting that a morphology intermediate to browsers and grazers may represent a fitness valley resulting from the inability to exploit both browse and graze efficiently. These results are important for future paleoecological studies and should be used as a guide for determining which dietary proxies are appropriate to the research question. J. Morphol., 2011.

Improving the repeatability of low magnification microwear methods using high dynamic range imaging

Abstract The recent advent of low magnification microwear analysis has allowed the efficient study of entire vertebrate faunas using only an optical stereomicroscope. Photographic visualization of microwear by this means has proven difficult, however, and, as a result, few high-resolution photos of low magnification microwear have been published. The repeatability of the method has also been questioned because low magnification microwear analysis involves the visual inspection of microwear features. We show that the use of high dynamic range imaging improves the visualization of microwear features in photographs and that using these photographs as a counting medium increases the repeatability of the method. We also show that counting from the photographs allows us to accurately classify ungulates as browsers, grazers, or mixed feeders.

Mean Annual Precipitation Explains Spatiotemporal Patterns of Cenozoic Mammal Beta Diversity and Latitudinal Diversity Gradients in North America

Spatial diversity patterns are thought to be driven by climate-mediated processes. However, temporal patterns of community composition remain poorly studied. We provide two complementary analyses of North American mammal diversity, using (i) a paleontological dataset (2077 localities with 2493 taxon occurrences) spanning 21 discrete subdivisions of the Cenozoic based on North American Land Mammal Ages (36 Ma – present), and (ii) climate space model predictions for 744 extant mammals under eight scenarios of future climate change. Spatial variation in fossil mammal community structure (β diversity) is highest at intermediate values of continental mean annual precipitation (MAP) estimated from paleosols (∼450 mm/year) and declines under both wetter and drier conditions, reflecting diversity patterns of modern mammals. Latitudinal gradients in community change (latitudinal turnover gradients, aka LTGs) increase in strength through the Cenozoic, but also show a cyclical pattern that is significantly explained by MAP. In general, LTGs are weakest when continental MAP is highest, similar to modern tropical ecosystems in which latitudinal diversity gradients are weak or undetectable. Projections under modeled climate change show no substantial change in β diversity or LTG strength for North American mammals. Our results suggest that similar climate-mediated mechanisms might drive spatial and temporal patterns of community composition in both fossil and extant mammals. We also provide empirical evidence that the ecological processes on which climate space models are based are insufficient for accurately forecasting long-term mammalian response to anthropogenic climate change and inclusion of historical parameters may be essential.

EARLY EVOLUTION OF SEXUAL DIMORPHISM AND POLYGYNY IN PINNIPEDIA

Sexual selection is one of the earliest areas of interest in evolutionary biology. And yet, the evolutionary history of sexually dimorphic traits remains poorly characterized for most vertebrate lineages. Here, we report on evidence for the early evolution of dimorphism within a model mammal group, the pinnipeds. Pinnipeds show a range of sexual dimorphism and mating systems that span the extremes of modern mammals, from monomorphic taxa with isolated and dispersed mating to extreme size dimorphism with highly ordered polygynous harem systems. In addition, the degree of dimorphism in pinnipeds is closely tied to mating system, with strongly dimorphic taxa always exhibiting a polygynous system, and more monomorphic taxa possessing weakly polygynous systems. We perform a comparative morphological description, and provide evidence of extreme sexual dimorphism (similar to sea lions), in the Miocene‐aged basal pinniped taxon Enaliarctos emlongi. Using a geometric morphometric approach and combining both modern and fossil taxa we show a close correlation between mating system and sex‐related cranial dimorphism, and also reconstruct the ancestral mating system of extant pinnipeds as highly polygynous. The results suggest that sexual dimorphism and extreme polygyny in pinnipeds arose by 27 Ma, in association with changing climatic conditions.

Anterior Dentary Shape as an Indicator of Diet in Ruminant Artiodactyls

ABSTRACT Several studies have quantified selective capability in artiodactyls using skeletal correlates such as the premaxillary shape index (PSI), incisor arcade breadth (IAB), incisor arcade curvature (DOC), and incisor width ratio (IWR). These metrics are limited in their applicability because they do not account for the potential importance of muzzle length. Herein we apply a new method for the quantification of muzzle shape (the dentary shape index [DSI]) that combines dentary length and width. We find that browsers possess the longest, narrowest muzzles, whereas the muzzles of grazers are comparatively short and wide. Discriminant function analysis demonstrates that DSI yields the highest rate of correct dietary classification of all muzzle shape proxies proposed to date. Generalized estimating equations and phylogenetically independent contrasts show that the shape of the anterior dentary (DSI) is significantly correlated with both diet and premaxillary shape (PSI) across the cetartiodactyl tree. We suggest that the match between premaxillary and anterior dentary shape is important in cropping but it is the shape of the lower incisor arcade that is under direct dietary selection. We conclude that the length and width of the anterior dentary are functional determinants of selectivity in ruminant artiodactyls and that these traits have potentially evolved in response to selection for efficient feeding as it is related to the different dietary requirements of browsers and grazers.

Complexity of Ruminant Masticatory Evolution

The evolution of robust jaws, hypsodont teeth, and large chewing muscles among grazing ruminants is a quintessential example of putative morphological adaptation. However, the degree of correlated evolution (i.e., to what extent the grazer feeding apparatus represents an evolutionary module), especially of soft and hard tissues, remains poorly understood. Recent generation of large datasets and phylogenetic information has made testing hypotheses of correlated evolution possible. We, therefore, test for correlated evolution among various traits of the ruminant masticatory apparatus including tooth crown height, jaw robustness, chewing muscle size, and characters of the molar occlusal surfaces, using phylogenetic and nonphylogenetic comparative methods as well as phylogenetic evolutionary model selection. We find that the large masseter muscles of grazing ruminants evolved with the inclusion of grass in the diet, an increase in the proportion of occlusal enamel bands oriented parallel to the chewing stroke, and possibly hypsodonty. We suggest that the masseter evolved under two evolutionary regimes: i) selection for higher masticatory forces during chewing and ii) flattening of the tooth profile, which resulted in reduced tooth guidance and, thus, a requirement for more chewing muscle activity during each chewing stroke, in agreement with previous research. The linear jaw metrics (depth of the mandibular angle, mandibular angle width, and length of the superficial masseteric scar) all show correlated evolution with hypsodonty and the proportion of enamel bands oriented parallel to the chewing stroke. We suggest that changes in the shape of the mandible represent the combined effects of selection for a reorientation of the chewing stroke, so as to emphasize horizontal translation of the teeth, and accommodation of high‐crowned teeth. Our analyses show that the ruminant feeding apparatus is an evolutionary mosaic with its various components showing both correlated and independent evolution. J. Morphol. 275:1093–1102, 2014.

Cycad (Cycadales) Chromosome Numbers Are Not Correlated with Genome Size

Premise of research. Change in base number of chromosomes per nucleus is usually believed to result from whole-genome duplication or from duplication or elimination of a single chromosome (aneuploidy). However, chromosome numbers can also change via mechanisms with no gain or loss of nuclear DNA, such as fusion or fission of chromosomes. Methodology. We previously determined amount of DNA per nucleus (2C values) using flow cytometry of leaf tissue. We tested whether Cycadales (cycad) chromosome numbers are correlated with these 2C values using ordinary least squares (OLS) and phylogenetic generalized least squares (PGLS) regression. Pivotal results. Regardless of branch length estimation technique and evolutionary model, OLS and PGLS yielded no statistically significant relationship between chromosome number and genome size in our analysis of 22 species, encompassing all 11 extant cycad genera. Conclusions. Within cycad genera, chromosomal fissions or fusions are much more likely than large additions or deletions of nuclear DNA.

THE USE OF GROSS DENTAL WEAR IN DIETARY STUDIES OF EXTINCT LAGOMORPHS

Abstract Studies of paleoenvironments have commonly focused on large mammalian herbivores such as ungulates. Many localities, however, have yielded large numbers of small mammalian herbivores, including lagomorphs and rodents. These fossils represent an untapped paleoecological resource. However, the fossils are often in the form of isolated teeth, and microwear analysis cannot be used due to taphonomic alteration. As a result, we use ungulate gross dental wear as a model. The dental wear features of extant western Canadian lagomorphs are identified and used to create dietary categories that can be applied to make predictions about the diets of extinct forms. The Horse Local Fauna of the Cypress Hills Formation of Saskatchewan has yielded approximately 2,500 fossil specimens, of which nearly 300 are lagomorphs. Two leporid species (rabbits and hares) are present in the Horse Local Fauna, Palaeolagus temnodon and Megalagus brachyodon. Qualitative analysis of the gross dental wear of the lagomorphs of the Horse Local Fauna indicates that M. brachyodon was mainly folivorous and P. temnodon was primarily frugivorous, suggesting that the contemporaneous ecosystem was tree dominated. Gross dental wear analysis allows the use of small herbivores and isolated teeth in paleoecological studies. Studying the diets of small herbivorous mammals will allow more nearly complete reconstructions of past environments and will become increasingly important as more detailed reconstructions are required by paleontologists.

Can latitudinal richness gradients be measured in the terrestrial fossil record

Abstract. Studying the deep-time origins of macroecological phenomena can help us to understand their long-term drivers. Given the considerable spatiotemporal bias of the terrestrial fossil record, it behooves us to understand how much biological information is lost. The aim of this study is to establish whether latitudinal diversity gradients are detectable in a biased terrestrial fossil record. I develop a simulated fossilization approach, weighting the probability of terrestrial mammal species appearing in the fossil record based on body size and geographic-range size; larger species with larger range sizes are more likely to enter the fossil record. I create simulated fossil localities from the modern North American mammal record. I vary the percentage of species successfully fossilized and estimate the magnitude of the latitudinal diversity gradient (slope of the richness gradient and degree of species turnover). I find that estimates of the latitudinal diversity gradient are sensitive to the loss of species with small body size and geographic-range sizes. In some cases, simulated fossil-record bias completely obliterates evidence of declining richness with latitude, a phenomenon that is not ameliorated by the application of nonparametric richness estimation. However, if the rate of preservation is medium (50% of species) to high (75% of species), the magnitude of the latitudinal diversity gradient can be reliably estimated. Similarly, changes in the diversity gradient estimates are largely explained by differences in the diversity—climate relationship among iterations, suggesting that these relationships may be measurable in the fossil record.