Range-wide sources of variation in reproductive rates of northern spotted owls.

We conducted a range-wide investigation of the dynamics of site-level reproductive rate of northern spotted owls using survey data from 11 study areas across the subspecies geographic range collected during 1993-2018. Our analytical approach accounted for imperfect detection of owl pairs and misclassification of successful reproduction (i.e., at least one young fledged) and contributed further insights into northern spotted owl population ecology and dynamics. Both nondetection and state misclassification were important, especially because factors affecting these sources of error also affected focal ecological parameters. Annual probabilities of site occupancy were greatest at sites with successful reproduction in the previous year and lowest for sites not occupied by a pair in the previous year. Site-specific occupancy transition probabilities declined over time and were negatively affected by barred owl presence. Overall, the site-specific probability of successful reproduction showed substantial year-to-year fluctuations and was similar for occupied sites that did or did not experience successful reproduction the previous year. Site-specific probabilities for successful reproduction were very small for sites that were unoccupied the previous year. Barred owl presence negatively affected the probability of successful reproduction by northern spotted owls in Washington and California, as predicted, but the effect in Oregon was mixed. The proportions of sites occupied by northern spotted owl pairs showed steep, near-monotonic declines over the study period, with all study areas showing the lowest observed levels of occupancy to date. If trends continue it is likely that northern spotted owls will become extirpated throughout large portions of their range in the coming decades.

Life and death in a dynamic environment: Invasive trout, floods, and intraspecific drivers of translocated populations.

Understanding the relative strengths of intrinsic and extrinsic factors regulating populations is a long-standing focus of ecology and critical to advancing conservation programs for imperiled species. Conservation could benefit from an increased understanding of factors influencing vital rates (somatic growth, recruitment, survival) in small, translocated populations, which is lacking owing to difficulties in long-term monitoring of rare species. Translocations, here defined as the transfer of wild-captured individuals from source populations to new habitats, are widely used for species conservation, but outcomes are often minimally monitored, and translocations that are monitored often fail. To improve our understanding of how translocated populations respond to environmental variation, we developed and tested hypotheses related to intrinsic (density dependent) and extrinsic (introduced rainbow trout Oncorhynchus mykiss, stream flow and temperature regime) causes of vital rate variation in endangered humpback chub (Gila cypha) populations translocated to Colorado River tributaries in the Grand Canyon (GC), USA. Using biannual recapture data from translocated populations over 10 years, we tested hypotheses related to seasonal somatic growth, and recruitment and population growth rates with linear mixed-effects models and temporal symmetry mark-recapture models. We combined data from recaptures and resights of dispersed fish (both physical captures and continuously recorded antenna detections) from throughout GC to test survival hypotheses, while accounting for site fidelity, using joint live-recapture/live-resight models. While recruitment only occurred in one site, which also drove population growth (relative to survival), evidence supported hypotheses related to density dependence in growth, survival, and recruitment, and somatic growth and recruitment were further limited by introduced trout. Mixed-effects models explained between 67% and 86% of the variation in somatic growth, which showed increased growth rates with greater flood-pulse frequency during monsoon season. Monthly survival was 0.56-0.99 and 0.80-0.99 in the two populations, with lower survival during periods of higher intraspecific abundance and low flood frequency. Our results suggest translocations can contribute toward the recovery of large-river fishes, but continued suppression of invasive fishes to enhance recruitment may be required to ensure population resilience. Furthermore, we demonstrate the importance of flooding to population demographics in food-depauperate, dynamic, invaded systems.

An occupancy-based quantification of the highly imperiled status of desert fishes of the southwestern United States.

Desert fishes are some of the most imperiled vertebrates worldwide due to their low economic worth and because they compete with humans for water. An ecological complex of fishes, 2 suckers (Catostomus latipinnis, Catostomus discobolus) and a chub (Gila robusta) (collectively managed as the so-called three species) are endemic to the U.S. Colorado River Basin, are affected by multiple stressors, and have allegedly declined dramatically. We built a series of occupancy models to determine relationships between trends in occupancy, local extinction, and local colonization rates, identify potential limiting factors, and evaluate the suitability of managing the 3 species collectively. For a historical period (1889-2011), top performing models (AICc) included a positive time trend in local extinction probability and a negative trend in local colonization probability. As flood frequency decreased post-development local extinction probability increased. By the end of the time series, 47% (95% CI 34-61) and 15% (95% CI 6-33) of sites remained occupied by the suckers and the chub, respectively, and models with the 2 species of sucker as one group and the chub as the other performed best. For a contemporary period (2001-2011), top performing (based on AICc ) models included peak annual discharge. As peak discharge increased, local extinction probability decreased and local colonization probability increased. For the contemporary period, results of models that split all 3 species into separate groups were similar to results of models that combined the 2 suckers but not the chub. Collectively, these results confirmed that declines in these fishes were strongly associated with water development and that relative to their historic distribution all 3 species have declined dramatically. Further, the chub was distinct in that it declined the most dramatically and therefore may need to be managed separately. Our modeling approach may be useful in other situations in which targeted data are sparse and conservation status and best management approach for multiple species are uncertain.

Una Cuantificación Basada en la Ocupación del Estado Altamente en Peligro de los Peces del Desierto del Suroeste de los Estados Unidos Resumen Los peces del desierto son unos de los vertebrados en mayor peligro a nivel mundial debido a su bajo valor económico y a que compiten por el agua con los humanos. Un complejo ecológico de peces, dos miembros de la familia Catostomidae (Catostomus latipinnis, C. discobolus) y un bagre (Gila robusta) (todos manejados colectivamente como las llamadas tres especies), endémico a la cuenca estadunidense del Río Colorado, está afectado por múltiples factores estresantes y supuestamente ha declinado dramáticamente. Construimos una serie de modelos de ocupación para determinar las relaciones entre las tendencias de ocupación, la extinción local y las tasas de colonización, para identificar los factores limitantes potenciales y para evaluar la idoneidad del manejo colectivo de las tres especies. Durante un periodo histórico (1889 - 2011), los modelos con el mejor desempeño (AICc) incluyeron una tendencia positiva de tiempo en la probabilidad de extinción local y una tendencia negativa en la probabilidad de colonización local. La probabilidad de extinción local incrementó conforme disminuyó la frecuencia de inundaciones pos-desarrollo. Al final de la serie de tiempo, el 47% (95% IC 34 - 61) y el 15% (95% IC 6 - 33) de los sitios permanecieron ocupados por los catostómidos y el bagre, respectivamente, y los modelos con las dos especies de catostómidos como un grupo y el bagre como otro tuvieron el mejor desempeño. Para un periodo contemporáneo (2001 - 2011), los modelos con el mejor desempeño (basados en AICc) incluyeron a la mayor descarga anual. La probabilidad de extinción local disminuyó y la probabilidad de colonización local incrementó conforme incrementó la mayor descarga. Para el periodo contemporáneo, los resultados de los modelos que dividieron a las tres especies en grupos separados fueron similares a los resultados de los modelos que combinaron a los dos catostómidos pero no al bagre. Colectivamente, estos resultados confirmaron que las declinaciones de estos peces estuvieron fuertemente asociadas con el desarrollo del agua y que, en relación a su distribución histórica, las tres especies han declinado dramáticamente. Además, el bagre se distinguió por declinar más dramáticamente y por lo tanto tal vez requiera de manejo por separado. Nuestra estrategia de modelado puede ser útil en otras situaciones en las que los datos de importancia son escasos y el estado de conservación y la mejor estrategia de manejo para múltiples especies son inciertos.

Evaluating impacts using a BACI design, ratios, and a Bayesian approach with a focus on restoration.

Before-after-control-impact (BACI) designs are an effective method to evaluate natural and human-induced perturbations on ecological variables when treatment sites cannot be randomly chosen. While effect sizes of interest can be tested with frequentist methods, using Bayesian Markov chain Monte Carlo (MCMC) sampling methods, probabilities of effect sizes, such as a ≥20 % increase in density after restoration, can be directly estimated. Although BACI and Bayesian methods are used widely for assessing natural and human-induced impacts for field experiments, the application of hierarchal Bayesian modeling with MCMC sampling to BACI designs is less common. Here, we combine these approaches and extend the typical presentation of results with an easy to interpret ratio, which provides an answer to the main study question-"How much impact did a management action or natural perturbation have?" As an example of this approach, we evaluate the impact of a restoration project, which implemented beaver dam analogs, on survival and density of juvenile steelhead. Results indicated the probabilities of a ≥30 % increase were high for survival and density after the dams were installed, 0.88 and 0.99, respectively, while probabilities for a higher increase of ≥50 % were variable, 0.17 and 0.82, respectively. This approach demonstrates a useful extension of Bayesian methods that can easily be generalized to other study designs from simple (e.g., single factor ANOVA, paired t test) to more complicated block designs (e.g., crossover, split-plot). This approach is valuable for estimating the probabilities of restoration impacts or other management actions.

Risk-sensitive allocation in seasonal dynamics of fat and protein reserves in a long-lived mammal.

Body reserves of numerous taxa follow seasonal rhythms that are a function of temporal patterns in food availability and life-history events; however, tests of the theory underlying the allocation of somatic reserves for long-lived organisms are rare, especially for free-ranging mammals. We evaluated the hypothesis that allocation of somatic reserves to survival (i.e., metabolic processes) and reproduction should be sensitive to current nutritional state relative to seasonal thresholds in those reserves. Our goal was to reveal the linkages between nutrition and life-history traits to understand how long-lived, iteroparous organisms balance the allocation of somatic reserves to reproduction, while retaining reserves as insurance for survival in unpredictable environments. Our evaluation was based on seasonal dynamics in fat (measured as ingesta-free body fat; IFBFat) and protein reserves (measured as ingesta-free, fat-free body mass; IFFFBMass) of 136 female mule deer (Odocoileus hemionus) over 8 years. Although mean changes in fat and protein reserves were positive over summer and negative over winter, accretion and catabolism of those reserves was not consistent among individuals. Over winter, both lipid and protein stores available in autumn were catabolized in proportion to their availability above a post-winter threshold (5·8% IFBFat, 33 kg IFFFBMass); however, lean body tissue was spared at the expense of lipid reserves. Female deer mostly synthesized lean body tissue over summer and committed post-winter fat reserves to reproduction relative to their availability above an autumn threshold (>8·6% IFBFat), which was lowered by 2·8 percentage points (pp) for each additional young recruited. Mothers reduced their autumn fat threshold to secure current reproductive investment and, thereby, endured a cost of reproduction at the expense of fat accumulation. Allocation of somatic reserves occurred in a risk-sensitive framework; females allocated reserves relative to their availability above seasonal thresholds. In contrast to current notions of summer accretion and winter catabolism of body reserves, some individuals deposited reserves over winter and catabolized reserves over summer, mainly because regulation of individual condition was state-dependent. Consequently, behaviour and life-history strategies may be as much a function of nutritional contributions of the previous season as of the current one.

Effects of gonadotropin-releasing hormone immunization on reproductive function and behavior in captive female Rocky Mountain elk (Cervus elaphus nelsoni).

Fertility control is a potential method for managing overabundant wildlife populations; however, current technology is limited by duration of treatment efficacy and unacceptable side effects. The objective of this study was to determine the efficacy of a single immunization with gonadotropin-releasing hormone (GnRH) vaccine to suppress reproductive function in pregnant female elk and to evaluate potential behavioral and pathological side effects of treatment. Eighteen captive adult female elk were randomly allocated to one of two experimental groups. Ten females were administered a conjugated and adjuvanted GnRH vaccine intramuscularly, and eight elk received an adjuvant sham vaccine without conjugated GnRH. We compared success of existing pregnancy, neonatal survival, subsequent fertility, reproductive behavior rates, and side effects of treatment between January 2006 and January 2010. The GnRH vaccination did not affect existing pregnancy or calf survival during the year that it was applied; however, it reduced the proportion of pregnant females for 3 yr. Male precopulatory behavior rates exhibited toward GnRH-vaccinated females tended to be greater than those directed at sham-vaccinated females during the second half of the breeding season, when GnRH vaccinates continued to be proceptive. Strong immune and inflammatory responses, including robust GnRH antibody concentrations in GnRH vaccinates, and sterile pyogranulomatous injection site abscesses in both groups, were consistent with vaccination. In conclusion, this GnRH vaccine resulted in prolonged, albeit reversible, impairment of fertility, and is associated with extended reproductive behaviors and partial suppression of hypothalamic-pituitary-gonadal axis function in captive female elk.

THE RELATIONSHIP BETWEEN HARVEST MANAGEMENT AND CHRONIC WASTING DISEASE PREVALENCE TRENDS IN WESTERN MULE DEER (ODOCOILEUS HEMIONUS) HERDS.

We analyzed retrospective data on harvest management practices and corresponding chronic wasting disease (CWD) prevalence trends in 36 western US and Canadian mule deer (Odocoileus hemionus) management units (units). Our analyses employed logistic regression and model selection, exploiting variation in practices within and among jurisdictions to examine relationships between harvest management and apparent prevalence (the proportion of positive animals among those sampled). Despite notable differences in hunting practices among jurisdictions, our meta-analysis of combined data revealed strong evidence that the amount of harvest was related to CWD prevalence trends among adult male mule deer in the 32 units where prevalence at the start of the analysis period was ≤5%. All competitive models included the number of male deer harvested or number of hunters 1-2 yr prior as an explanatory variable, with increasing harvest leading to lower prevalence among males harvested in the following year. Competitive models also included harvest timing. Although less definitive than the number harvested, median harvest dates falling closer to breeding seasons were associated with lower prevalence in the following year. Our findings suggest harvest-when sufficient and sustained-can be an effective tool for attenuating CWD prevalence in adult male mule deer across western ranges, especially early in the course of an epidemic. Evidence of a broad relationship between the amount of harvest and subsequent changes in CWD prevalence among adult male mule deer provides an empirical basis for undertaking adaptive disease management experimentation aimed at suppressing or curtailing CWD epidemics.

Mountain lions prey selectively on prion-infected mule deer.

The possibility that predators choose prey selectively based on age or condition has been suggested but rarely tested. We examined whether mountain lions (Puma concolor) selectively prey upon mule deer (Odocoileus hemionus) infected with chronic wasting disease, a prion disease. We located kill sites of mountain lions in the northern Front Range of Colorado, USA, and compared disease prevalence among lion-killed adult (> or =2 years old) deer with prevalence among sympatric deer taken by hunters in the vicinity of kill sites. Hunter-killed female deer were less likely to be infected than males (odds ratios (OR) = 0.2, 95% confidence intervals (CI) = 0.1-0.6; p = 0.015). However, both female (OR = 8.5, 95% CI = 2.3-30.9) and male deer (OR = 3.2, 95% CI = 1-10) killed by a mountain lion were more likely to be infected than same-sex deer killed in the vicinity by a hunter (p < 0.001), suggesting that mountain lions in this area actively selected prion-infected individuals when targeting adult mule deer as prey items.

EFFECT OF ORAL COPPER SUPPLEMENTATION ON SUSCEPTIBILITY IN WHITE-TAILED DEER (ODOCOILEUS VIRGINIANUS) TO CHRONIC WASTING DISEASE.

Chronic wasting disease (CWD) is an infectious disease, but reported associations suggest several metals-especially copper (Cu) and manganese-potentially play a role in this and other prion diseases. To assess the utility of dietary Cu supplementation in protecting white-tailed deer (Odocoileus virginianus) from CWD, we compared incidence and disease course among individuals naturally exposed to CWD while being maintained on sustained-release Cu boluses or unsupplemented (control). Oral Cu supplementation increased liver tissue Cu concentrations compared to controls but did not affect susceptibility to CWD or survival after natural exposure in the captive white-tailed deer we studied. Over the 27 mo study, 89% (8/9) of the Cu-supplemented deer and 86% (6/7) of control deer became CWD-infected. Survival to 27 mo postexposure did not differ between Cu-supplemented and control deer: model-averaged survival probabilities to 27 mo were 0.45-0.47 for all combinations of Cu treatment and PRNP gene haplotype presence. The PRNP gene haplotype influenced the probability of deer remaining biopsy negative for at least 17 mo but did not affect overall susceptibility.

Methods to quantify variable importance: implications for the analysis of noisy ecological data.

Determining the importance of independent variables is of practical relevance to ecologists and managers concerned with allocating limited resources to the management of natural systems. Although techniques that identify explanatory variables having the largest influence on the response variable are needed to design management actions effectively, the use of various indices to evaluate variable importance is poorly understood. Using Monte Carlo simulations, we compared six different indices commonly used to evaluate variable importance; zero-order correlations, partial correlations, semipartial correlations, standardized regression coefficients, Akaike weights, and independent effects. We simulated four scenarios to evaluate the indices under progressively more complex circumstances that included correlation between explanatory variables, as well as a spurious variable that was correlated with other explanatory variables, but not with the dependent variable. No index performed perfectly under all circumstances, but partial correlations and Akaike weights performed poorly in all cases. Zero-order correlations was the only measure that detected the presence of a spurious variable, whereas only independent effects assigned overlap areas correctly once the spurious variable was removed. We therefore recommend using zero-order correlations to eliminate predictor variables with correlations near zero, followed by the use of independent effects to assign overlap areas and rank variable importance.

Recolonizing wolves and mesopredator suppression of coyotes: impacts on pronghorn population dynamics.

Food web theory predicts that the loss of large carnivores may contribute to elevated predation rates and, hence, declining prey populations, through the process of mesopredator release. However, opportunities to test predictions of the mesopredator release hypothesis are rare, and the extent to which changes in predation rates influence prey population dynamics may not be clear due to a lack of demographic information on the prey population of interest. We utilized spatial and seasonal heterogeneity in wolf distribution and abundance to evaluate whether mesopredator release of coyotes (Canis latrans), resulting from the extirpation of wolves (Canis lupus) throughout much of the United States, contributes to high rates of neonatal mortality in ungulates. To test this hypothesis, we contrasted causes of mortality and survival rates of pronghorn (Antilocapra americana) neonates captured at wolf-free and wolf-abundant sites in western Wyoming, USA, between 2002 and 2004. We then used these data to parameterize stochastic population models to heuristically assess the impact of wolves on pronghorn population dynamics due to changes in neonatal survival. Coyote predation was the primary cause of mortality at all sites, but mortality due to coyotes was 34% lower in areas utilized by wolves (P < 0.001). Based on simulation modeling, the realized population growth rate was 0.92 based on fawn survival in the absence of wolves, and 1.06 at sites utilized by wolves. Thus, wolf restoration is predicted to shift the trajectory of the pronghorn population from a declining to an increasing trend. Our results suggest that reintroductions of large carnivores may influence biodiversity through effects on prey populations mediated by mesopredator suppression. In addition, our approach, which combines empirical data on the population of interest with information from other data sources, demonstrates the utility of using simulation modeling to more fully evaluate ecological theories by moving beyond estimating changes in vital rates to analyses of population-level impacts.

A meta-BACI approach for evaluating management intervention on chronic wasting disease in mule deer.

Advances in acquiring and analyzing the spatial attributes of data have greatly enhanced the potential utility of wildlife disease surveillance data for addressing problems of ecological or economic importance. We present an approach for using wildlife disease surveillance data to identify areas for (or of) intervention, to spatially delineate paired treatment and control areas, and then to analyze these nonrandomly selected sites in a meta-analysis framework via before-after-control impact (BACI) estimates of effect size. We apply these methods to evaluate the effectiveness of attempts to reduce chronic wasting disease (CWD) prevalence through intensive localized culling of mule deer (Odocoileus hemionus) in north-central Colorado, USA. Areas where surveillance data revealed high prevalence or case clusters were targeted by state wildlife management agency personnel for focal scale (on average <17 km2) culling, primarily via agency sharpshooters. Each area of sustained culling that we could also identify as unique by cluster analysis was considered a potential treatment area. Treatment areas, along with spatially paired control areas that we constructed post hoc in a case-control design (collectively called "management evaluation sites"), were then delineated using home range estimators. Using meta-BACI analysis of CWD prevalence data for all management evaluation sites, the mean effect size (change of prevalence on treatment areas minus change in prevalence on their paired control areas) was 0.03 (SE = 0.03); mean effect size on treatment areas was not greater than on paired control areas. Excluding cull samples from prevalence estimates or allowing for an equal or greater two-year lag in system responses to management did not change this outcome. We concluded that management benefits were not evident, although whether this represented true ineffectiveness or was a result of lack of data or insufficient duration of treatment could not be discerned. Based on our observations, we offer recommendations for designing a management experiment with 80% power to detect a 0.10 drop in prevalence over a 6-12-year period.

Epidemiology of chronic wasting disease in free-ranging mule deer: spatial, temporal, and demographic influences on observed prevalence patterns.

We analyzed chronic wasting disease (CWD) prevalence data from mule deer populations in northcentral Colorado, USA, to discern the likely influences of temporal, spatial, and demographic factors on patterns observed in naturally infected populations. In addition to reaffirming spatial heterogeneity among wintering mule deer subpopulations, we report marked differences in CWD prevalence by sex and age groups as well as clear local trends of increasing prevalence over a 7-yr period. Prevalence of CWD differed by age (yearling vs. adult), sex, and geographic area at two different spatial scales (game management unit or population unit winter range) and increased over time at both geographic scales. Disease status (positive or negative) was not independent of age for males (n=285, df=6, chi2=18.4, P=0.005) or females (n=387, df=8, chi2=17.2, P=0.028). Among males, prevalence increased and then declined across age classes, peaking in 5- to 6-yr-old individuals; among females, prevalence showed no definite age-related pattern. Demographic, spatial, and temporal factors all appear to contribute to the marked heterogeneity in CWD prevalence in endemic portions of northcentral Colorado, USA. These factors likely combine in various ways to influence epidemic dynamics on both local and broad geographic scales.

Relative vulnerability of chronic wasting disease infected mule deer to vehicle collisions.

We estimated chronic wasting disease (CWD) prevalence among vehicle-killed mule deer (Odocoileus hemionus) in select data analysis units (DAUs) in northern Colorado, USA, and compared these with estimated CWD prevalence among mule deer of the same sex sampled in the vicinity of collision sites to assess relative vulnerability of CWD-infected individuals to vehicle collisions. Twenty-five of 171 vehicle-killed mule deer tested positive for CWD (overall prevalence=0.146, 95% confidence interval [CI]=0.097-0.208); 173 of 2,317 deer sampled in the vicinity of these vehicle-killed deer tested positive (overall prevalence=0.075, 95% CI=0.064-0.085). In nine of ten DAU x sex comparisons, relative risk of CWD infection tended to be higher among vehicle-killed deer (range of estimated relative risks=1.6-15.9). Spongiform encephalopathy was detected in 12 of 20 (60%; 95% CI=39-81%) CWD-positive deer killed by vehicles and in 79 of 180 (44%; 95% CI=37-52%) CWD-positive deer detected via random sampling (relative risk=1.37; 95% CI=0.92-2.03), suggesting that infected deer killed by vehicles tended to be in later stages of disease than those killed by hunters. Our data offer evidence that CWD-infected mule deer may be relatively vulnerable to vehicle collisions. It follows that sampling of vehicle-killed mule deer may be exploited to increase efficiency of surveillance programs designed to detect new foci of CWD infection; moreover, evidence of increased susceptibility to vehicle collisions may aid in understanding vulnerability of CWD-infected individuals to other forms of death, particularly predation.

Select tissue mineral concentrations and chronic wasting disease status in mule deer from North-central Colorado.

Trace mineral imbalances have been suggested as having a causative or contributory role in chronic wasting disease (CWD), a prion disease of several North American cervid species. To begin exploring relationships between tissue mineral concentrations and CWD in natural systems, we measured liver tissue concentrations of copper, manganese, and molybdenum in samples from 447 apparently healthy, adult (> or = 2 yr old) mule deer (Odocoileus hemionus) culled or vehicle killed from free-ranging populations in north-central Colorado, United States, where CWD occurs naturally; we also measured copper concentrations in brain-stem (medulla oblongata at the obex) tissue from 181 of these deer. Analyses revealed a wide range of concentrations of all three minerals among sampled deer (copper: 5.6-331 ppm in liver, 1.5-31.9 ppm in obex; manganese: 0.1-21.4 ppm in liver; molybdenum: 0.5-4.0 ppm in liver). Bayesian multiple regression analysis revealed a negative association between obex copper (-0.097; 95% credible interval -0.192 to -0.006) and the probability of sampled deer also being infected with CWD, as well as a positive association between liver manganese (0.158; 95% credible interval 0.066 to 0.253) and probability of infection. We could not discern whether the tendencies toward lower brain-stem copper concentrations or higher systemic manganese concentrations in infected deer preceded prion infection or rather were the result of infection and its subsequent effects, although the distribution of trace mineral concentrations in infected deer seemed more suggestive of the latter.

The role of geographic information systems in wildlife landscape epidemiology: models of chronic wasting disease in Colorado mule deer.

The authors present findings from two landscape epidemiology studies of chronic wasting disease (CWD) in northern Colorado mule deer (Odocoileus hemionus). First, the effects of human land use on disease prevalence were explored by formulating a set of models estimating CWD prevalence in relation to differences in human land use, sex and geographic location. Prevalence was higher in developed areas and among male deer suggesting that anthropogenic influences (changes in land use), differences in exposure risk between sexes and landscape-scaled heterogeneity are associated with CWD prevalence. The second study focused on identifying scales of mule deer movement and mixing that had the greatest influence on the spatial pattern of CWD in north-central Colorado. The authors hypothesised that three scales of mixing - individual, winter subpopulation and summer subpopulation - might control spatial variation in disease prevalence. A fully Bayesian hierarchical model was developed to compare the strength of evidence for each mixing scale. Strong evidence was found indicating that the finest mixing scale corresponded best to the observed spatial distribution of CWD prevalence. This analysis demonstrates how information on the scales of spatial processes that generate observed patterns can be used to gain insight into the epidemiology of wildlife diseases when process data are sparse or unavailable.

Low frequency of PrP genotype 225SF among free-ranging mule deer (Odocoileus hemionus) with chronic wasting disease.

The prion protein (PrP) gene was characterized in 1482 free-ranging mule deer (Odocoileus hemionus) from Wyoming and Colorado. Using DNA sequences from 363 deer, dimorphisms at codons 20 (aspartate/glycine) and 225 [serine (S)/phenylalanine (F)] were found; silent changes occurred at codons 131 (tyrosine) and 247 (isoleucine). The remaining samples were surveyed for codon 225 genotype and all were characterized for chronic wasting disease (CWD) infection status. A total of 112 deer with the genotype 225SF or FF were found, of which one was CWD-positive; 1370 were 225SS, with 289 positive for CWD. Among CWD-negative deer, the frequency of 225SF/FF genotypes was 9.3 % but among CWD-positive deer it was only 0.3 %. For all samples combined, CWD status was not independent of codon 225 genotype (P<0.0001). The odds that a deer of the 225SS genotype was CWD-infected were 30 times greater (95 % confidence intervals=4-213) than for a 225SF deer. The proportion of 225SF animals in sampled subpopulations varied from 0 to 18 %; the CWD prevalence varied from 0 to 25 %. However, no relationship was observed between genotype frequency and CWD prevalence in different areas. The PrP sequences of experimentally infected mule deer were analysed from pre-existing projects and 10 animals were found with 225SF genotypes, all of which were positive for CWD. Data available from some of these animals suggest that the 225SF genotype could be associated with longer incubation periods in CWD infection compared with the 225SS genotype.