The independent evolution of the sexes may often be constrained if male and female homologous traits share a similar genetic architecture. Thus, cross-sex genetic covariance is assumed to play a key role in the evolution of sexual dimorphism (SD) with consequent impacts on sexual selection, population dynamics and the speciation process. I used quantitative genetics tools to assess the importance of sex-specific genetic variance in facilitating the evolution of body mass and horn size SD in wild bighorn sheep from Ram Mountain, Alberta. I also developed a bighorn sheep genetic linkage map composed of 247 microsatellite markers to gain insights about the genetic architecture of trait variation. Finally, I conducted systematic reviews and meta-analyses of published cross-sex genetic correlations (rMF, a standardized estimate of cross-sex genetic covariance) to test basic hypotheses about the importance of sex-specific genetic variance in the evolution of SD and mechanisms responsible for generating such variance. My results demonstrated that sex-specific genetic variance was present in bighorn sheep and that it likely played an important role in alleviating intralocus sexual conflicts. The quantitative trait locus (QTL) mapping analysis resulted in the identification of numerous loci influencing body mass and horn dimensions, some of which had apparent sex-specific effects. An analysis of 553 rMF estimates recovered from 114 published sources allowed demonstrating that 1) the evolution of SD was generally constrained by positive cross-sex genetic covariance, 2) levels of SD were often sub-optimal, and 3) sex-specific genetic variance was an important mechanism allowing the evolution of SD. In addition, I confirmed the long-standing hypothesis of a general decline in rMF with age. Sexual dimorphism is an important evolutionary phenomenon, but our understanding of its evolution is still limited. After decades of speculation, my research has provided clear empirical evidence for the importance of sex-specific genetic variance in allowing its evolution. / Ecology

Human encroachment into wilderness areas can influence the persistence of wildlife populations by decreasing and degrading habitat, displacement, and decreasing survival. For bighorn sheep (Ovis canadensis), some human activities are detrimental, causing both physiological stress and habitat abandonment. Between 1979 and 2000, human recreation has increased over 300% in areas occupied by desert bighorn sheep (O. c. nelsonii) in southeastern Utah. We investigated if an increase in human activity in areas used by bighorns affected their behavior. We observed 34 bighorn sheep using focal-animal sampling for >14 hrs to compare time spent grazing and scanning between areas of high and low human use. We identified group size, presence or absence of a lamb, distance to escape terrain, and human use (high versus low) as potential explanatory variables that influenced grazing and scanning times, and created an a priori list of models based on these variables. We used Akaike's Information Criterion adjusted for small sample sizes (AICc) to rank models, and used model selection to find a best approximating model (lowest AICc value) for both behaviors. Desert bighorn sheep spent less time grazing and more time scanning in high human use areas (22% grazing, 29% scanning) than in low human use areas (54% grazing, 8% scanning). Caution should be taken when considering which areas or trails should be opened during these important seasons to minimize and reduce additional stresses to bighorns caused by human activity. Bighorn sheep populations experienced significant declines after European settlement in North America. Today, the primary practice of bighorn sheep conservation is through population restoration and augmentation from remnant source populations. We conducted a 9-year telemetry study for a source population of desert bighorn sheep in Canyonlands National Park, Utah. We captured and collared 58 bighorn sheep from 2002-2009. To estimate annual and seasonal survival, we used known-fate analysis in Program MARK 4.1. We used model selection to test hypotheses for bighorn survival, including sex, age, human use, year, and month, as possible explanatory variables. There were 20 mortalities during the study. Annual survival ranged from 83% - 88% with no significant variation among any of the years. Model selection results showed that the top six models included a temporal variable (e.g. season or month), and carried 92% of the AICc weight. Population persistence for bighorn sheep can be compromised by high levels of predation, habitat fragmentation, and disease transmitted from domestic sheep. We suggest that land managers continue to maintain the separation of domestic sheep from bighorns in CNP. We also recommend that survival studies continue to ensure that future translocation projects do not occur at the expense of the source population.

This dissertation presents an investigation and design optimization of energy absorbent protective systems that protect the brain. Specifically, the energy absorption characteristics of the bighorn sheep skull-horn system were quantified and used to inform a topology optimization performed on a football helmet facemask leading to reduced values of brain injury indicators. The horn keratin of a bighorn sheep was experimentally characterized in different stress states, strain rates, and moisture contents. Horn keratin demonstrated a clear strain rate dependence in both tension and compression. As the strain rate increased, the flow stress increased. Also, increased moisture content decreased the strength and increased ductility. The hydrated horn keratin energy absorption increased at high strain rates when compared to quasi-static data. The keratin experimental data was then used to inform constitutive models employed in the simulation of bighorn sheep head impacts at 5.5 m/s. Accelerations values as high as 607 G’s were observed in finite element simulations for rams butting their heads, which is an order of magnitude higher than predicted brain injury threshold values. In the most extreme case, maximum tensile pressure and maximum shear strains in the ram brain were 245 kPa and 0.28, respectively. These values could serve as true injury metrics for human head impacts. Finally, a helmeted human head Finite Element (FE) model is created, validated, and used to recreate impacts from a linear impactor. The results from these simulations are used to train a surrogate model, which is in turn utilized in multi-objective design optimization. Brain injury indicators were significantly reduced by performing multi-objective design optimization on a football helmet facemask. In particular, the tensile pressure and maximum shear strain in the brain decreased 7.5 % and 39.5 %, respectively when comparing the optimal designs to the baseline design. While the maximum tensile pressure and maximum shear strain values in the brain for helmeted head impacts (30.2 kPa and 0.011) were far less than the ram impacts (245 kPa and 0.28), helmet impacts up to 12.3 m/s have been recorded, and could easily surpass these thresholds.

Many wildlife populations globally are experiencing unprecedented declines, and without accurate and precise estimates of abundance, we will not be able to conserve these vulnerable species. Remote cameras have rapidly advanced as wildlife monitoring tools and may provide accurate and precise estimates of abundance that improve upon traditional methods. Using remote cameras to estimate abundance may be less expensive, less intrusive, less dangerous, and less time consuming than other methods. While it is apparent that remote cameras have a place in the future of wildlife monitoring, research, and management, many questions remain concerning the proper use of these tools. In an effort to answer some of these questions, we used remote cameras to study a population of Rocky Mountain bighorn sheep (Ovis canadensis) in Utah, USA from 2012 to 2014. In Chapter 1, we compared methods using remote cameras against 2 traditional methods of estimating abundance. In Chapter 2, we evaluated the relationship between deployment time of cameras and proportion of photos needed to be analyzed to obtain precise estimates of abundance. We found that methods using remote cameras compared favorably to traditional methods of estimating abundance, and provided a number of valuable advantages. In addition, we found that remote cameras can produce precise estimates of abundance in a relatively short sampling period. Finally, we identified the optimal sampling period to produce precise estimates of abundance for our study population. Our findings can help researchers better utilize the potential of remote cameras, making them a more suitable alternative to traditional wildlife monitoring.

Many wildlife populations globally are experiencing unprecedented declines, and without accurate and precise estimates of abundance, we will not be able to conserve these vulnerable species. Remote cameras have rapidly advanced as wildlife monitoring tools and may provide accurate and precise estimates of abundance that improve upon traditional methods. Using remote cameras to estimate abundance may be less expensive, less intrusive, less dangerous, and less time consuming than other methods. While it is apparent that remote cameras have a place in the future of wildlife monitoring, research, and management, many questions remain concerning the proper use of these tools. In an effort to answer some of these questions, we used remote cameras to study a population of Rocky Mountain bighorn sheep (Ovis canadensis) in Utah, USA from 2012 to 2014. In Chapter 1, we compared methods using remote cameras against 2 traditional methods of estimating abundance. In Chapter 2, we evaluated the relationship between deployment time of cameras and proportion of photos needed to be analyzed to obtain precise estimates of abundance. We found that methods using remote cameras compared favorably to traditional methods of estimating abundance, and provided a number of valuable advantages. In addition, we found that remote cameras can produce precise estimates of abundance in a relatively short sampling period. Finally, we identified the optimal sampling period to produce precise estimates of abundance for our study population. Our findings can help researchers better utilize the potential of remote cameras, making them a more suitable alternative to traditional wildlife monitoring.

A recently emerging engineering design approach entails studying the brilliant design solutions found in nature with an aim to develop design strategies that mimic the remarkable efficiency found in biological systems. This novel engineering approach is referred to as bio-inspired design. In this context, the present study quantifies the structure-property relations in bighorn sheep (Ovis canadensis) horn keratin, qualitatively characterizes the effects of a tapered spiral geometry (the same form as in a ram’s horn) on pressure wave and impulse mitigation, describes the stress attenuation capabilities and features of a ram’s head, and compares the structures and mechanical properties of some energy absorbent natural materials. The results and ideas presented herein can be used in the development of lightweight, energy absorbent, bio-inspired material designs. Among the most notable conclusions garnered from this research include: Horn keratin behaves in an anisotropic manner similar to a long fiber composite. Moisture content dominates the material behavior of horn keratin more than anisotropy, age, and stress-state. This makes moisture content the most influential parameter on the mechanical behavior of horn keratin. Tapered geometries mitigate the impulse generated by a stress wave due to the convergent boundary and a continually decreasing cross sectional area such that greater uniaxial stresses and subsequent axial deformation arises. Furthermore, the tapered geometry introduces small shear stresses that further decrease the impulse. Spiral geometries attenuate the impulse generated by a stress wave by the introduction of shear stresses along the length of the spiral. These shear stresses introduce transverse displacements that function to lessen the impulse. When both a taper and spiral geometry are used in a design, their synergistic effects multiplicatively reduce the impulse Tough natural materials have a high porosity, which makes them light-weight, while increasing their compressive energy absorption ability. Biomaterials whose functions include protection and energy absorption feature a multiscale, hierarchical, composite structure. The constituent materials are arranged in such ways to achieve a synergistic effect, where the properties of the composite exceed the properties of its constituents. Biological materials are therefore not confined to the law of mixtures.