Seasonal Habitat Use and Movements of Mountain Goats, Oreamnos americanus, in East-central British Columbia

To identify the potential for adverse effects of forest development on Mountain Goats (Oreamnos americanus), we documented the patterns of forest use by goats and the factors influencing goat habitat use. We used a combination of 15 very high frequency (VHF) and six global positioning system (GPS) radiocollars to document the distribution and movements of 21 (15 female, 6 male) goats from 1997 to 1999 in the mountains surrounding the Robson Valley in east-central British Columbia. Because canopy closure reduces the likelihood that a GPS receiver will obtain a location fix, we estimated that GPS collars underrepresented forest use by about 23%. Three goats used separate winter and summer ranges separated by 8–13 km, while most simply exhibited seasonal shifts in elevation. In winter, goats were more often at lower elevations, in commercial forest stands, on southerly aspects, and moved less each hour and over the course of the winter. Goat use declined in areas >500 m from escape terrain and goats were found lower in elevation from evening to dawn compared to daylight hours. Collared goats used high elevation licks, which were either within their home range, or in two cases, 6 and 14 km from their typical home range. We documented use of known mid-elevation mineral licks by three collared goats, but no use of known low elevation (valley bottom and lower slopes) mineral licks. Robson Valley goats appeared to be at relatively low risk from disturbances related to logging, because although forest use was documented during winter, it occurred primarily on high elevation, steep slopes where trees are currently of low commercial value, and goats made little use of low elevation mineral licks. We recommend that in this area a forested buffer of 500 m around cliffs be left to reduce the possibility of adverse effects on goats especially, on southerly aspects above 1300 m

Alpine ungulate movement: Quantification of spatiotemporal environmental energetics and social interaction

Species movement, an animal’s ability to change its location, is a fundamental property of life, and animals have diverse physical and behavioural attributes that are believed to enhance efficient travel and optimization of resources. Quantifying movement energetics and returns to examine these ideas over relevant time- and space scales is, however, problematic. In this thesis, I set out to develop and use advanced biologging tag technology to determine a second by second account of the behaviour and location of tagged animals to unveil where and when key behaviours are occurring, to answer key questions about feeding and social behaviour, allocation in space and the energetic costs associated with different movement decisions. Specifically, I used accelerometers, magnetometers, temperature and pressure sensors with GPS units in animal-attached loggers to examine key questions linking movement, energetics and feeding and aggressive behaviours in 3 wild- and 3 domestic ungulate species in mountainous landscapes in the French Alps, monitored for periods between 30 and 200 days. To obtain high-frequency data using electronic devices for long periods, I had to first design new housings to attach safely the loggers to the animals and develop methods for weather proofing the loggers. I designed, using CAD-designa and 3D printing, different housing types and used ‘Guronic’ resin to shockproof and waterproof circuit boards. This allowed me to obtain logging data for up to 200 days. To give a location per second but stay within ethical weight restrictions, the dead-reckoning method to reconstruct fine-scale movements between low resolution GPS fixes was adopted. To improve the accuracy of dead-reckoning estimates I improved the method using behavioural definition to identify real moves (steps, grazing, moving) and distinguish it from resting, grooming and other behaviours not leading to a displacement of the animal in space, allowing to selectively filter data to be dead-reckon. Using the data collected, I showed that central-place-based, but free-roaming, domestic goats exhibited efficient space-use by having time-dependent fanning out from their central place, which reduced local resource depletion. Models predicted that area-use increased logarithmically with herd size and duration. These finding could lead to improved livestock management in multi-functional alpine landscapes, to reduce the risk of over-grazing and manage interactions with other grazing species and clonflicts with other landuse needs. The goat grazing patterns were compared to those of wild ibex and revealed goats to be more adaptable, with the ibex being particularly vulnerable to changes in temperature, exacerbated by them preferring steep slopes with associated high metabolic costs and heat generation during ascent. These results could further inform management decisions regarding the survival of alpine ibex under projected climate change. Furthermore I developed new biologging approaches to investigate social interactions, specifically head-clashing in both species. This agonistic behaviour was associated with competition and the rut in ibex and was quantified using methods first developed for the domestic goat, where the behaviour appeared to relate primarily to competition for food. Using the goat as a surrogate species, the behaviour could be identified and mapped for the ibex, which highlighted areas and times important for head-clashing, including drastic increases during the rut. Finally, movement data and proxies for energy expenditure from three domestic species (sheep, cows and goats) and three wild species (ibex, mouflon and chamois) was utilised to produce species-specific energy landscapes across the terrains they used. This indicated that different anatomies and behaviours resulted in different, species-specific, movement costs for specific topographies and habitats. Energy use for travel across heterogeneous space depends, therefore, on the species concerned. These findings thus highlight the importance to consider that species with different life histories and ecological needs use landscapes in contrasting ways and my results can provide a more refined evidence base for the management and conservation of these species in alpine grasslands. These biologging approaches allow now also to address further management issues such as the responses to disturbances from tourists (hiking, skiers, etc.) and even reveal how species are more susceptible to climate change

Evaluating population estimates of mountain goats based on citizen science

Citizen science programs that use trained volunteers may be a cost-effective method for monitoring wildlife at large spatial and temporal scales. However, few studies have compared inferences made from data collected by volunteers to professionally collected data. In Glacier National Park (GNP), Montana, I assessed whether citizen science is a useful method to monitor mountain goat (Oreamnos americanus) populations. I compared estimates of mountain goat abundance by volunteers at 32 sites throughout GNP to estimates by biologists and raw counts from aerial surveys at a subset of 25 and 11 sites, respectively. I used multiple observer surveys to calibrate the indices of abundance for the effect of observer variation between volunteers and biologists. I used N-mixture models, which calculated detection probability through patterns of detection and non-detection to obtain estimates of abundance. Population estimates made by citizen science overlapped estimates by biologists and estimates from previous research. Density estimates from aerial surveys were lower, possibly due to imperfect detection during aerial surveys or due to violation of the assumption of population closure. Mean detection probability from multiple observer surveys for biologists was significantly higher and less variable than that of volunteers, but was not a suitable correction factor, because it was not consistent across all densities of mountain goats. Volunteer experience did not significantly influence detection probability or abundance estimates. Abundance estimates by volunteers were influenced by number of site visits. More frequent site visits balanced out lower detection probability by volunteers and resulted in abundance estimates that were less variable than those of biologists. When large spatial and temporal coverage can be achieved, citizen science can provide mountain goat population estimates that are statistically similar to those of biologists. However, neither estimates by volunteers or biologists had sufficient statistical power to detect a 30% decline in mountain goat population size over 10 years. Power by volunteers could be increased by reducing the number of sites and increasing surveys/site or by continuing monitoring over a longer time frame (i.e., 30 years). Citizen science programs can contribute to long term monitoring when properly designed

Utilizing Remote Sensing and Geospatial Techniques to Determine Detection Probabilities of Large Mammals

Whether a species is rare and requires protection or is overabundant and needs control, an accurate estimate of population size is essential for the development of conservation plans and management goals. Wildlife censuses in remote locations or over extensive areas are logistically difficult, frequently biased, and time consuming. My dissertation examined various techniques to determine the probability of detecting animals using remotely sensed imagery. We investigated four procedures that integrated unsupervised classification, texture characteristics, spectral enhancements, and image differencing to identify and count animals in remotely sensed imagery. The semi-automated processes had relatively high errors of over-counting (i.e., greater than 60%) in contrast to low (i.e. less than 19%) under-counting errors. The single-day image differencing had over-counting errors of 53% while the manual interpretation had over-counting errors of 19%. The probability of detection indicates the ability of a process or analyst to detect animals in an image or during an aerial wildlife survey and can adjust total counts to estimate the size of a population. The probabilities of detecting an animal in remotely sensed imagery with semi-automated techniques, single-day image differencing, or manual interpretation were high (e.g. ≥ 80%). Single-day image differencing resulted in the highest probability of detection suggesting this method could provide a new technique for managers to estimate animal populations, especially in open, grassland habitats. Remotely sensed imagery can be successfully used to identify and count animals in isolated or remote areas and improve management decisions. Sightability models, used to estimate population abundances, are derived from count data and the probability of detecting an animal during a census. Global positioning systems (GPS) radio-collared bison in the Henry Mountains of south-central Utah provided a unique opportunity to examine remotely sensed physiographic and survey characteristics for known occurrences of double-counted and missed animals. Bison status (detected, missed, or double-counted) was determined by intersecting helicopter survey paths with bison travel paths during annual helicopter surveys. The probability of detecting GPS-collared bison during the survey ranged from 91% in 2011 to 88% in 2012

Habitat modeling using path analysis: delineating mountain goat habitat in the Washington Cascades

A 70-90% decline in mountain goat (Oreamnos americanus) populations in Washington State over the past few decades has spurred the need for an improved understanding of seasonal goat-habitat relationships. Habitat use data have been collected from 46 radio-collared mountain goats across their native range in Washington State. Using Geographical Information Systems (GIS), I explored relationships between use and availability of habitat. To overcome issues of autocorrelation, I compared actual mountain goat paths with available paths of matched identical spatial topology and used multi-scale path analysis to explore various ecologically informed relationships between landscape structure and the movements of mountain goats at the home range scale. I extracted used and available (randomized) paths at 4 scales of analysis using square extraction windows of 0.06, 4.4, 15.2, and 56.2 ha that were centered on each point along the path. Matched case logistic regression allowed me to determine the spatially and temporally explicit scales that were the strongest predictors of seasonal and year-round mountain goat habitat from a suite of predictor variables. I found that for year-round habitat, mountain goats chose both abiotic and biotic components of their landscape including; parkland, areas of high solar loading, terrain that is rugged, and terrain that allows escape from predators. This analysis represents one of the most extensive landscape-level habitat relationship studies conducted on mountain goats. Additionally, my methodological approach is applicable to other species-habitat association analyses

Cougar Predation Rates and Prey Composition in the Pryor Mountains of Wyoming and Montana

Understanding predator-prey dynamics is a fundamental principle of ecology and an ideal component for management decisions. Across North America, the impact of cougars (Puma concolor) on their prey varies regionally. To document the relationships between cougars, bighorn sheep (Ovis canadensis canadensis), mule deer (Odocoileus hemionus), and feral horses (Equus caballus) on the Bighorn Canyon National Recreation Area and the Pryor Mountains, we deployed GPS collars on 6 cougars (the total number residing on the study area), and visited their clusters to determine predation rates and foraging patterns. We examined the composition of cougar kills by species, mule deer sex-age classes, prey size classes, season, and cougar sex. As a measure of selection, we examined the composition of prey killed relative to the composition of ungulates obtained during an aerial survey. We found mule deer were the primary prey, while bighorn sheep constituted secondary prey. While cougars selected for bighorn sheep, this was attributable to a single cougar. Among mule deer, female cougars killed more does and male cougars killed more bucks. Family groups had the highest predation rates (i.e., the shortest time intervals between kills), while adult males had the lowest rate. During the study, cougars were not depredating any feral horses in the area. Maintaining predator and prey numbers will require agencies to monitor and manage all fauna within this complex ecosystem. Habitat manipulations may be necessary to increase populations of deer and bighorn sheep, while continued management of feral horses will be required to reduce competition with native ungulates

Resource selection and abundance estimation of moose: Implications for caribou recovery in a human altered landscape

Woodland caribou (Rangifer tarandus caribou) are threatened across Canada due to human disturbance altering predator-prey dynamics. The niche specialization of caribou enables them to survive in nutrient-poor habitats spatially separated from other ungulates and their shared predators. The conversion of old-growth forests to young seral stands is hypothesized to increase the abundance of moose (Alces alces), the dominant prey for wolves (Canis lupus), resulting in apparent competition. We first examined habitat selection of moose in 2 regions with differing intensities of human disturbance in west-central Alberta and east-central British Columbia to assess how human disturbance affects the spatial separation of moose and caribou. We built resource selection functions with data from global positioning system (GPS) collars deployed on 17 moose (8 in a region with high and 9 in a region with low human disturbance) at 2 spatial scales. Our results indicated that moose in our study area make forage-risk tradeoffs in a hierarchical fashion similar to caribou, potentially eroding spatial separation in human disturbed landscapes. We also evaluated the spatial partitioning of resources by comparing resource use with GPS locations from 17 moose and 17 paired caribou using logistic regression. As expected, human disturbance decreased the resource partitioning between moose and caribou. Thus, systematic moose management and monitoring will be essential for caribou conservation. Currently, a Stratified Random Block (SRB) survey design is widely used to estimate moose populations, but these surveys are expensive and often result in imprecise population estimates when not corrected for sightability bias. We evaluated the application of distance sampling as an alternative to SRB surveys, especially for use in caribou ranges. To correct for moose missed on the transect line, where a detection rate of 100% is critical, we developed a sightability model using 21 radio-collared moose. After correcting for sightability, distance sampling was more precise and efficient than SRB surveys. In this way, more efficient distance sampling methodology can be an important tool for caribou conservation. Combined, our results showed the importance of moose management in caribou ranges due to decreased spatial separation between both ungulate species in disturbed landscapes

GPS tracking in high mountain landscapes: insights into the movement ecology of female alpine ibex (Capra ibex ibex L. 1758).

The three studies reported in this thesis have been conducted on the Alpine ibex population of the Marmolada-Monzoni, in the north-eastearn Italian Alps. A summary for each study is given below. Chapter I: Land morphology, season and individual activity influence GPS fix acquisition rates and location error in an alpine ungulate. The use of GPS technologies in wildlife research has greatly increased the opportunities for addressing ecological issues that affect ultimately the conservation of the species. However, in order to formulate accurate and unbiased conclusions in studies of movement ecology with GPS-tracking systems, it is necessary to understand the sources of potential bias and error associated with this technology, under specific environmental conditions and taking into account the behavioural patterns of the species monitored. In chapter I, I first present the results of a field trial with stationary collars scheduled to attempt a location fix every 30 min during 24 hours cycles. The collars were positioned in 64 locations throughout the study area in order to sample different land cover categories and topographic conditions. GPS collars performances were influenced mainly by available sky view. When sky view was higher than 70%, acquired as respect to scheduled locations were close to 100%, and location accuracy was within 10 m for 75% of acquired locations. When sky view was below 70 %, the proportion of acquired locations dropped to 75% and location error increased to within 20 m for 75% of locations acquired. I then examined a database of more than 90.000 attempted locations from 11 GPS-tagged females of Alpine ibex to assess the temporal trends in fix acquisition rate, and how it was influenced by habitat features of daily areas used, by individual activity, and by climate and weather variables. I found that fix acquisition rate was very good and scarcely variable in summer, but could drop to less than 85% during the coldest months and at night in winter. Fix acquisition rate was strongly and positively influenced by individual activity and declined, especially in winter, in periods of adverse weather and lower than average temperatures. Most probably female ibex, when inactive and seeking for shelter, use microhabitats providing cover that obstructs the satellites signal, so reducing fix acquisition rate. I concluded that, although with an adequate screening procedure for identifying outliers the accuracy of locations received from different habitat conditions may remain good, the acquired locations underestimate the use of habitats providing shelter and the periods of adverse weather. In general, the results underline the importance of combining stationary tests with tests on free-ranging animals when assessing GPS bias and accuracy in field conditions. Chapter II: Determinants of home range size across spatio-temporal scales in a high mountain ungulate. The high seasonality of Alpine habitats might have strong effects on the spatial strategies of large herbivores. In the second chapter, I obtained a database of 672 estimates of weekly home ranges (232 in summer and 440 in winter) and 160 estimates of monthly home ranges (64 in summer and 96 in winter) from 15 female ibexes, and analysed it to describe intra-annual patterns of spatial use and to asses how it was influenced by climate, food resources and individual conditions. I used the k-LoCoH method to calculate the areas used at two spatial scales: the home range (HR; calculated on 95% of locations) and the core area (CA, calculated on 50% of the locations). At all temporal and spatial scales, the areas used by females were very small in deep winter, progressively increased until a peak in mid-summer, and then dropped again. This pattern was very marked, with a 15-20-fold increase in size from the winter minimum to the summer maximum. The HR and CA size was positively correlated with daylight, but was more synchronized with indexes of climate and vegetation phenology, as absolute temperature and average NDVI of the study area. After having defined biologically meaningful seasons with a clustering approach based on step distances and habitat features associated with locations, I then analysed, within seasons and correcting for temporal trends, the effects of the stochastic variability of climatic and weather conditions and of food resources on the size of ranges used. I found that, in winter, HRs and CAs at all temporal scales decreased strongly when snow was deeper, or precipitations more abundant, while in summer they decreased with increasing food resources (indexed by the average NDVI value or proportion of vegetation in the HRs or CAs). Also slope, which I used as an index of refuge areas from predators but also of snow accumulation, had a marked negative effect on the size of areas used. In contrast, individual conditions, as age class and reproductive status, did not influence with consistent patterns the spatial behaviour of females. These results highlight the peculiar strategy of spatial use of female ibex, which appear to be extremely energy conservative in winter and aimed at optimizing the use of food resources in summer. The understanding of factors driving spatial behaviour of female ibex is fundamental to conserve key wintering areas and habitats, and to predict how future climate changes might impact on the species. Chapter III: Validation of a non-invasive technique for estimating diet quality in an alpine ungulate. In seasonal environments, the behavioural patterns of large herbivores are shaped by the availability of forage resources, which affect the individual performance and reproduction success. Faecal nitrogen (FN) and Faecal Neutral Detergent Fibre (FNDF) have been proposed as indicators of diet quality in wildlife species. In the study reported in chapter III, I aimed at evaluating the use of faecal N, and secondarily of faceal NDF, to describe patterns of diet quality in Alpine ibex. Since chemical analyses are costly and time consuming, I also verificed whether NIRS estimates of faecal N might provide results as accurate as those of chemical analyses. From late June to November, I collected fresh samples of female and male ibex faeces, which were analyzed for FN and FNDF using Chemical analysis and three different NIRS instruments with variable wavelength ranges and approach (reflectance or transmittance). In order to verify possible relationships, I also associated to each sample the NDVI index of greenness of a surrounding buffer area. NIRS analyses gave good predictions for N, and only slightly lower for NDF, provided that the instrument used operated over a wide spectral range and in reflectance. Faecal N decreased, and FNDF increased, with Julian date, suggesting a reduction in diet quality thorugh summer and autumn. Females tended to have higher FN and lower FNDF contents than males, suggesting the ability to select a diet of better quality. These patterns were best described by data from chemical analyses, but were closely approximated by those from the best NIRS method. The NDVI of the buffer area surrounding faecal samples did not influence indexes of diet quality. I concluded that FN estimated with NIRS techniques could be a useful tool for studying patterns of diet quality in Alpine ibex. The declining FN and increasing FNDF values from summer to autumn suggest that ibex do not have the ability to contrast, with alternative food sources or with increasing selectivity, the decline in vegetation quality. This emphasizes the importance of energy-saving strategies during the winter, and of exploiting the short availability of good food resources in spring

Evaluating Methods for Pathogen Spillover Detection and Forecasting in Desert Bighorn Sheep (\u3cem\u3eOvis canadensis nelsoni \u3c/em\u3e)

Managers mitigate the effects of disease on wildlife populations by detecting outbreaks and predicting disease transmission. Bighorn sheep (Ovis canadensis) populations face threat from the introduced pathogen Mycoplasma ovipneumoniae (M. ovipneumoniae). In the eastern Mojave Desert, managers detect pathogen outbreaks in desert bighorn (O. c. nelsoni) through aerial population surveys, which produce irregular and coarse population information. Furthermore, managers model the risk of M. ovipneumoniae transmission amongst desert bighorn populations, however these models assume that desert bighorn space use is unchanging across years. Here, I evaluate methods used by managers to detect disease outbreaks and an assumption for predicting disease transmission. The results will provide managers guidance for passive vital rate monitoring and refine models informing managers of transmission risk. In my second chapter, I evaluated camera traps’ ability to represent population metrics indicative of disease outbreak and persistence. I compared camera- and in personderived bighorn counts to quantify how well cameras detect presence, group sizes, and lamb:ewe ratios. Across camera trap sites and years examined, the probability that camera traps detected present bighorn sheep ranged from 44% to 100%, with per-minute probability ranging from 6% to 84%. The probability that camera traps detected the arrival of new groups ranged from 18% to 100%. Camera-derived group size estimates exhibited substantial negative bias across all sites and demographic groups, and bias varied across group sizes present. Across all sites, the multiplicative change between camera and observer lamb:ewe ratios ranged from 1.79 to 2.46. In my third chapter, I evaluated the assumption that desert bighorn space use is consistent across years when environmental condition varies. I did this by examining how home range sizes varied as a function of dynamic environmental condition, including environmental moisture and plant biomass, and static landscape characteristics, including slope, vector ruggedness measure (VRM), and distance to water. I found that ram home range sizes responded more to temporally dynamic variables than ewe home ranges, while ewe home range sizes responded more to landscape features. These results suggest that transmission risk models should incorporate bighorn space use changes across years