Ruffed grouse populations are low in Appalachian forests, possibly because low habitat quality negatively affects survival, condition, and reproduction. Through the Appalachian Cooperative Grouse Research Project (ACGRP) researchers tracked >1500 radioed grouse at 10 study sites (1996__2002). To improve our understanding of Appalachian grouse habitat ecology, I carried out two primary analyses of this database. First, grouse should be under selective pressure to minimize movements, so I studied factors associated with variation in home range size. Second, importance of a habitat is affected by an individual's resource needs, and I investigated factors associated with variation in selection of "preferred" habitats. Both approaches yielded important insights into the species' regional habitat ecology. As elsewhere, clearcuts, which afford escape cover, formed the cornerstone of grouse habitat in the region. However, a number of other factors were also important. At the root of this was a divergence in habitat ecology between grouse inhabiting the two major forest types in the region. In oak-hickory forests nutritional constraint strongly influenced habitat use. Grouse home ranges increased 2.5x following poor hard mast crops, and at these times grouse increased use of alternate foraging habitats. Grouse, especially females and broods, made extensive use of mesic bottomlands and forest edges, which in oak-hickory forests support relatively abundant soft mast and herbaceous forages. In contrast, grouse inhabiting mixed mesophytic forests were insensitive to hard mast, did not select bottomlands, reduced use of forest edges, and increased use of clearcuts. I feel that greater abundance of birch, cherry, and aspen, buds of which are a high quality winter food, relieves nutritional stress on grouse inhabiting mesophytic forests. A general inference was that grouse attempted to balance competing strategies of maximizing either survival or condition, and the expression of this tradeoff was mediated by forest composition. Also presented here were studies of radiotelemetry error, roost site selection, and suitability of prescribed burning as a habitat improvement technique. In the closing chapter I make recommendations for managing Appalachian forests for grouse, which focus on improving winter foraging habitat, brood habitat, and escape cover, all of which are limiting in Appalachian forests. / Ph. D.

State-space modelling is a powerful tool to study ecological systems. The direct inclusion of uncertainty, unification of models and data, and ability to model unobserved, hidden states increases our knowledge about the environment and provides new ecological insights. I extend the state-space framework to create multi-species models, showing that the ability to model ecosystem interactions is limited only by data availability. State-space models are fit using both Bayesian and Frequentist methods, making them independent of a statistical school of thought. Bayesian approaches can have the advantage in their ability to account for missing data and fit hierarchical structures and models with many parameters to limited data; often the case in ecological studies. I have taken a Bayesian model fitting approach in this thesis. The predator-prey interactions between the hen harrier (Circus cyaneus) and red grouse (Lagopus lagopus scoticus) are used to demonstrate state-space modelling’s capabilities. The harrier data are believed to be known without error, while missing data make the cyclic dynamics of the grouse harder to model. The grouse-harrier interactions are modelled in a multi-species state-space model, rather than including one species as a covariate in the other’s model. Finally, models are included for the harriers’ alternate prey. The single- and multi-species state-space models for the predator-prey interactions provide insight into the species’ management. The models investigate aspects of the species’ behaviour, from the mechanisms behind grouse cycles to what motivates harrier immigration. The inferences drawn from these models are applicable to management, suggesting actions to halt grouse cycles or mitigate the grouse-harrier conflict. Overall, the multi-species models suggest that two popular ideas for grouse-harrier management, diversionary feeding and habitat manipulation to reduce alternate prey densities, will not have the desired effect, and in the case of reducing prey densities, may even increase the harriers’ impact on grouse chicks.

The capercaillie is the largest member of the grouse family extant in Scotland. This species is reported to have become extinct during the 1700s. It is also reported to have been reintroduced to Scotland from Sweden during the 1800s. There have been many assertions made about the underlying causes of the decline of the species; however the specific causal factors remain unknown. The reintroduction of the capercaillie to Scotland in the 1800s is the only successful reintroduction of a grouse species ever to have occurred in the world. The specific factors behind the success of the reintroduction also remain unknown. This thesis examines the extent to which a selection of historical documentary evidence can help to establish both the causes of the 18th century decline of the capercaillie in Scotland and the successes of the 19th century reintroduction. The methodology of this thesis incorporates facets from the fields of both environmental science and history. The methodology includes three steps. The first step involves the selection of a series of potential critical factors that may have been responsible for the decline of the species in the 1700s; these critical factors were selected from the present day understanding of the ecology and the behaviour of the capercaillie. The second step of the methodology includes the surveying of a series of historical documentary sources. From these surveys historical observations of the species were gathered. The historical documentary sources selected for examination in this thesis include what are referred to here as ‘primary historical source material’ and ‘secondary historical source material’. The majority of the primary historical source material was gathered from the hand written manuscripts of the Breadalbane estate, held at the National Archives of Scotland (NAS) and the Atholl estate held at the Blair Castle Charter Room in Blair Atholl. Other select primary historical source material consulted to a lesser extent, due to time constraints, includes the Forfeited Estates (1745) Inventory and the Baron Court Records for Strathspey and Urquhart (1617–1683) from the Grant estate muniments; both held at the NAS. The secondary historical source material was gathered from published and edited literary collections that include historical accounts of the species. The third and final step of the methodology involves the synthesis of both the historical and environmental information in order to establish to what extent the causes of both the decline of the species in the 1700s and success of its reintroduction in the 1800s can be realised. The findings from this thesis assert that the capercaillie was resident in Scotland from, at least, the end of the Medieval. Moving forward from the Middle Ages this thesis presents observations of the capercaillie throughout the historical period. These observations of the capercaillie appear in many different historical accounts. In some instances these observations are fleeting and do not form the main subject of the particular document in question. In other instances accounts of the species are much more detailed and include references to the ecology and behaviour of the bird. The level of detail included in an observation aside, the frequency with which the species is referred to in the sample of historical documents suggests that sections of the Scottish human population were familiar with the species, in various locations and at various times throughout history. By the 17th century the capercaillie is reported as beginning to become rare in some locations while still remaining comparatively abundant in others. The number of instances where the species is referred to as becoming rare in the historical documents increases between the 17th and the 19th centuries. Despite the reported scarceness of the species in Scotland from around the 17th century onwards, the capercaillie is recorded as persisting in Scotland until around the end of the 1700s. By the early 1800s the number of observations of the species in the secondary historical source material increases. All of the observations in the secondary sources from the early 1800s record the absence of the species from localities and regions of Scotland. No new evidence was found in either the primary or secondary historical source material to challenge the supposition that the capercaillie did become extinct in Scotland after 1785. No detailed quantitative data was available for analysis of the decline of the species. Thus, to investigate the extent to which the historical accounts can help explain the specific causal factors of the reported decline, a synthesis of the environmental and historical data was necessary. The findings of this synthesis suggest that the naturally occurring Scottish population of capercaillie probably persisted in the form of a metapopulation. The two hundred years between the 17th and 19th century most likely saw the extinction of capercaillie sub-populations, before the loss of the overall population of capercaillie around 1785. The sample historical documentary evidence alludes to this pattern of local and/or regional extinction of sub-populations. The cause or causes of the extinction of these sub-populations has focussed on five limiting or critical factors known to affect the species today. These five factors are climate change, particularly weather effects associated with the Little Ice Age, habitat loss and deterioration, disturbance, human hunting and predation by species other than humans also contributed to the species’ extinction. The extent to which these critical factors affected each sub-population would have varied between regions of Scotland occupied by the capercaillie in history. This thesis proposes that there was no single or combination of specific critical factors that were ultimately responsible for the decline of the capercaillie in Scotland during the 1700s. In some areas the capercaillie sub-populations would have most likely died out as a result of habitat loss and deterioration and climate change. Whereas in others predation and inbreeding may have been the critical factors responsible for the species’ demise. More detailed information referring to the capercaillie was found in the historical documentary source material for the period post-extinction (i.e. 1800 onwards). Contrary to popular understanding numerous attempts to reintroduce the capercaillie to Scotland were carried out before the Marquis of Breadalbane’s successful programme in 1837. The historical documentary evidence reports early attempts to reintroduce the species to locations such as the Isle of Arran in 1807, on the Duke of Atholl’s estate in 1822 and on the Earl of Mar’s estate in 1824. None of these reintroduction programmes are reported to have been successful in establishing a ‘wild’ population. However, in some instances the captive rearing programmes initiated did bear some fruit and captive reared birds were sent from Dunkeld by the Duke of Atholl to Kenmore and were used in Breadalbane’s successful reintroduction in 1837. The historical documents report two causes for the failure of these early reintroduction attempts. The first is the sudden death of captive birds, most likely as a result of choking due to stress as observed in recent rearing programmes (i.e. Moss 1986). The reintroduced Arran population is reported to have become extinct in this fashion. The second reported cause of failure is predation by species other than man. For example the entire population of birds brought to Scotland by the Earl of Mar were predated when released on his estate. This thesis offers two critical factors as explanations for the remarkable success of the capercaillie’s reintroduction to Scotland in the 19th century. The first is the method by which the reintroduction was carried out; specifically, the re-establishment of a series of capercaillie sub-populations in different regions of Scotland.

Greater sage-grouse (Centrocercus urophasianus; hereafter, sage-grouse) are a species of conservation concern in the rangelands of western North America due to their dramatic decline over the last half century. Effective conservation and management of sensitive species requires an understanding of how species respond to management actions. We examined two aspects of the reproductive phases of sage-grouse: nest predation, and habitat selection by female sage-grouse with chicks. In Chapter 1, we developed resource selection functions to assess the influence of mechanical treatments of mountain big sagebrush (Artemisia tridentata vaseyana) on habitat selection by greater sage-grouse with chicks. Post-treatment sage-grouse showed stronger selection for treatments and treatment edges than did pre-treatment sage-grouse. This altered pattern of selection by sage-grouse with broods suggests mechanical treatments may be a suitable way to increase use of mountain big sagebrush during the brooding period. In Chapter 2, we assessed the effect of habitat edges on nest predation of sage-grouse. The "edge effect" hypothesis states that habitat edges are associated with reduced nest success for birds. We tested the edge effect hypothesis using 155 nest locations from 114 sage-grouse. We derived edge metrics for 11 habitat cover types to determine which variables may have affected nest predation. We found support for the edge effect hypothesis in that nest predation increased with increasing edge density of paved roads. We provide evidence that the edge effect hypothesis may apply to greater sage-grouse and their habitats. Based on our results, we recommend minimizing disturbances that fragment critical nesting habitat of greater sage-grouse.

My research provided new information concerning the management, ecology, and conservation of greater sage-grouse (Centrocercus urophasianus). I report the results of an experiment using strategic intensive sheep grazing to enhance the quality of greater sage-grouse brood-rearing habitat. Although forb cover, an important component of brood-rearing habitat, responded positively to the grazing treatment, the response of other habitat variables was suppressed because the plots were not protected from domestic and wild herbivores during the years following the treatments. Measurements taken in grazing exclosures confirmed that herbivory by both large and small animals had significant impacts on vegetation. However, despite the suppressed habitat response, sage-grouse preferred the treated plots over the controls. In another chapter, I modeled survival rates of sage-grouse chicks to 42-days of age. Average chick survival across my study was high (39%). Survival varied across years and was affected by demographic, behavioral, and habitat factors. The top habitat model indicated that chick survival was positively related to grass cover and was higher in areas dominated by black sagebrush (Artemisia nova) than in big sagebrush (A. tridentata). The top model with demographic/behavioral factors indicated that survival was affected by interactions between hen age and brood mixing as well as between hatch date and brood mixing. In my last chapter I report on a survey of Utah sage-grouse hunter motivations and satisfaction. In 2008 and 2009 I surveyed over 600 sage-grouse hunters in Utah to determine why they chose to apply for sage-grouse hunting permits and what factors contributed to a satisfactory hunting experience. Originally, I had hypothesized that the impending Endangered Species Act listing petition for greater sage-grouse motivated hunters to pursue the species before they lost the opportunity. This hypothesis was not supported by the data. The majority of hunters indicated that they chose to hunt sage-grouse because it was a tradition or because it provided an opportunity to spend time outdoors with family. Additionally, Utah sage-grouse hunter satisfaction was influenced by whether or not the hunter was successful in harvesting at least one bird.

Declining populations of greater sage-grouse (Centrocercus urophasianus; hereafter sage-grouse) have increased stakeholder concerns regarding the management and stability of the species range-wide. Numerous conservation strategies have been identified to restoring sage-grouse population declines to include species translocations. Translocations have been used for many different wildlife species to help sustain genetic heterogeneity, reestablish, and augment declining populations. In a recent translocation study, researchers identified the protocols used to successfully translocate sage-grouse to restore declining populations in Strawberry Valley, Utah. This translocation occurred in a high elevation basin buffered by geomorphic barriers. I evaluated these protocols for use in translocating sage-grouse to augment a declining population that inhabited Anthro Mountain in northwest Utah. Anthro Mountain is a high elevation mountain dominated by sagebrush (Artemisia spp.) void of geomorphic barriers. I compared annual production, survival (i.e., vital rates), habitat use, and movements of translocated birds and their progeny to the resident population. Lastly, I described the integration of translocated birds with resident birds and the overall efficacy of the translocation effort. I radio-collared and monitored 60 translocated female sage-grouse from Parker Mountain, Utah over a 2-year period (2009 and 2010) and compared their vital rates to 19 radio-marked resident sage-grouse. Adult survival was similar for resident and translocated birds, but higher for both groups in 2010 than in 2009. However, overall survival of both resident and translocated birds was lower than range-wide survival estimates. Nest success was slightly higher for resident birds than translocated birds but positively correlated to grass height for both groups. Chick survival was also slightly higher for resident birds than for translocated birds, and higher overall in 2010 than in 2009. Chick survival was positively correlated to grass cover for both groups. Translocated birds used similar habitats and exhibited migration behaviors similar to resident birds. From a methodology perspective, the translocations protocols were successful because the translocated birds quickly acclimated to the release area, and their survival and reproductive success were similar to the resident birds. The effect of the translocation on augmenting the local population was inconclusive.

Dans le contexte du changement global et de la perte de la biodiversité, la prise en compte de cette dernière dans la gestion forestière reste encore très générale. Les démarches ne sont pas encore opérationnelles alors que les gestionnaires ont besoin d'outils destinés directement à la planification. Ainsi, il est devenu urgent d'améliorer les approches d'évaluation et de suivi de la biodiversité et de développer un cadre conceptuel à partir des indicateurs biologiques qui puisse servir dans la prise des décisions. Ce travail se repose sur une approche associant l'enjeu de conservation d'une espèce sauvage patrimoniale dans un paysage de montagne à la maîtrise d'une gestion forestière qui détermine la qualité de son habitat. Dans le cadre concret de la gestion forestière d'un Parc Naturel Régional où les enjeux sont multiples (écologiques, économiques, touristiques), les acteurs concernés s'interrogent moins sur le « comment gérer » que sur le « où gérer » ; en d'autres termes, définir les périmètres de bonne qualité d'habitat afin d'optimiser la gestion et à l'échelle optimale. Notre travail est moins de procurer des outils aux gestionnaires que de poser les jalons scientifiques d'une approche holistique inspirée de l'écologie du paysage. Nous partons ainsi de l'hypothèse selon laquelle les variables environnementales ainsi que la structure du paysage influencent la présence d'une espèce indicatrice de la diversité biologique d'un milieu. Dans ce travail nous nous appuyons sur cette hypothèse pour développer une démarche spatialement explicite qui permettrait d'atteindre l'objectif de conservation dans le cadre d'un processus de gestion forestière adaptative, en étudiant le cas de la gélinotte des bois dans le Parc Naturel Régional de Grande Chartreuse. Ce modèle prédictif de l'occurrence de la gélinotte des bois en Chartreuse s'appuie sur des indicateurs environnementaux et paysagers aisément mesurables et spatialisables. Ceci permettra d'établir un diagnostic de la qualité de l'habitat, d'approfondir les connaissances sur les relations entre la structure du paysage et l'espèce à différentes échelles et de déterminer les secteurs d'habitat qui sont les plus appropriés à la gélinotte des bois en zone de montagne. La représentation spatiale de l'hétérogénéité à différentes échelles permet d'envisager de nombreuses applications dans les domaines de la conservation et de la gestion forestière multifonctionnelle. Nos résultats, relevant d'une approche spatialement explicite, valident l'hypothèse selon laquelle la qualité de l'habitat de la gélinotte peut être modélisée avec une précision acceptable uniquement à partir d'indices paysagers, s'ils sont calculés sur la bonne étendue. Ils constituent donc une avancée pour une aide à la gestion, puisqu'ils préfigurent d'un outil permettant la localisation de zones prioritaires sur lesquelles la conservation de la gélinotte peut être optimisée. / In the context of global change and current loss of biodiversity, there is an urgent need to improve spatial approaches for assessing and monitoring biodiversity in order to develop a conceptual framework based on biological indicators that can be used in conservation planning. The problem still remains in terms of decisions and knowledge on where to set appropriate conservation targets. Hence, we need detailed and reliable information about forest landscape structure and composition and methods for estimating this information over the whole spatial domain. This work is based on an approach involving conservation of a patrimonial species in a mountain landscape. The Regional Natural Park in the French sub-Alps, used for this study encounters multiple conflicts (i.e. ecological, economic, tourism pressures). The stakeholders are then very concern on "where to manage" in other words, they need tools to define the perimeter of good quality habitat to optimize management at an optimal scale. Landscape metrics are widely used to explore the spatial heterogeneity of landscapes that influence several species distribution. Few empirical researches have comparatively examined their indicator value for certain species occurrence at different scales. Within the framework of this study we test hypothesis concerning the best scale at which Hazel Grouse respond to the landscape structure. Our aim was to evaluate significant relations between different landscape level indicators and species occurrence and to assess the effects of the extent of the considered landscape on their performance. Results showed that spatial scale affected the performance of the metrics, since Hazel Grouse occurrence was usually better predicted at relatively smaller extents of surrounding landscape. In all, outputs of this work constitute support for forest management, as they provide a methodological framework and indicators for prioritizing areas in terms of habitat quality where species conservation planning can be optimized.

My research was focused on greater sage-grouse (Centrocercus urophasianus; hereafter "sage-grouse") nest-site selection, nest success, and hen survival in relation to avian predators. The trade-off between using habitat and avoiding predators is a common decision for prey species including sage-grouse. In Chapter 2, I compared avian predator densities at sage-grouse nest and brood locations to random locations. Sage-grouse were located where densities of small, medium, and large avian predators were 65-68% less than random locations. The effects of anthropogenic and landscape features on habitat use of sage-grouse hens have not been evaluated relative to avian predator densities. In Chapter 3, I compared anthropogenic and landscape features and avian predator densities among sage-grouse locations (nest, early-brood, late-brood) and random locations. I found sage-grouse hens chose locations with lower avian predator densities compared to random locations, and selected locations farther away from anthropogenic and landscape features. Depredation of sage-grouse nests can be an influential factor limiting their productivity. Predator removal has been simultaneously proposed and criticized as a potential mitigation measure for low reproductive rates of sage-grouse. In Chapter 4, I hypothesized that sage-grouse nest success would be greater in areas where Wildlife Services lowered common raven (Corvus corax: hereafter "raven") density. I found that Wildlife Services decreased raven density by 61% during 2008-2011 but I did not detect a direct improvement to sage-grouse nest success. However, sage-grouse nest success was 22% when ravens were detected within 550 m of a sage-grouse nest and 41% when no raven was detected within 550 m. In Chapter 5, I assessed interactive effects of corvid densities relative to anthropogenic and landscape features on sage-grouse nest success. I found that sage-grouse nest success was positively correlated with rugged habitat. Survival of breeding-age birds is the most important demographic parameter driving sage-grouse abundance. In Chapter 6, I evaluated the effect of raptor densities, proximity to anthropogenic and landscape features, and hen behavior on survival of sage-grouse hens. I found that sage-grouse hen survival was negatively correlated with golden eagle (Aquila chrysaetos) density, proximity to anthropogenic and landscape features, and hen parental investment (nesting and brood-rearing).

Snowshoe hares (Lepus americanus) are a critically important prey species for Canada lynx (Lynx canadensis). Determination of snowshoe hare distribution and abundance is needed in western Wyoming for lynx conservation. We used linear regression to examine the correlations between snowshoe hare density, as determined by mark-recapture estimates, and fecal pellet plot counts on both uncleared and annually cleared plots on the Bridger-Teton National Forest, western Wyoming. We found significant correlations between hare density estimates and fecal pellet counts for both uncleared and annually cleared pellet counts; however the relationship was much stronger for annually cleared pellet counts. Adjusting the buffer size by omitting hard habitat edges (not used by hares) around the trapping grids improved correlations between hare density and fecal pellet counts further. We recommend pellet counts from annually cleared plots be used when precise estimates of snowshoe hare abundance are required. Though precision is sacrificed when using uncleared plots, they are useful as a coarse index of habitat use by hares. The derived regression equations should be used to identify foraging habitat for lynx in western Wyoming. In addition to snowshoe hares, in western Wyoming red squirrels (Tamiasciurus hudsonicus) and grouse (Bonasa umbellus and Dendragapus obscurus) are used by Canada lynx. Whether young forests or older multi-storied forests contain more snowshoe hares, red squirrels, and grouse in western Wyoming is currently unknown. We estimated snowshoe hare density, and indexed red squirrel and forest grouse abundance in 3 classes of 30-70-year-old lodgepole pine (Pinus contorta) and 4 classes of mature multi-storied forest with a spruce (Picea engelmannii)-fir (Abies lasiocarpa) component. We recorded landscape and forest structure characteristics to understand how these influence lynx prey abundance. Overall, snowshoe hares, red squirrels, and forest grouse were more abundant in multi-storied forests than young forests. Forest attributes that predicted prey abundance were often more prevalent in multi-storied forests. Results from this study suggest that multi-storied forests with a spruce-fir component were disproportionately important to snowshoe hares, red squirrels, and forest grouse in western Wyoming. Canada lynx conservation efforts should focus on maintaining, enhancing, and promoting multi-storied forests in this region.

The greater sage-grouse (Centrocercus urophasianus; sage-grouse) is a species of conservation concern in Utah and range-wide due to declines in populations and threats to sagebrush habitat on which they depend. To effectively conserve the species, detailed site-specific knowledge of ecology and distribution is needed. To expand knowledge of local populations within the West Box Elder Sage Grouse Management Area (SGMA) and gain insights into the effectiveness of vegetation treatments intended to benefit sagegrouse, I radio marked and tracked 123 (68 female, 55 male) sage-grouse and conducted sage-grouse pellet surveys on 19 conifer removal projects. Widespread habitat restoration measures designed to benefit sage-grouse have highlighted the need for prioritization tools to optimize placement of sage-grouse habitat projects. I generated seasonal habitat models to predict sage-grouse habitat use within the West Box Elder SGMA using a suite of vegetation and topographical predictors and known sage-grouse locations. Model fit was good with brood, early summer, late summer, lekking (early spring), and non-breeding models reporting an AUC of >0.90; nest and winter models reported an AUC of 0.87 and 0.85, respectively. A vegetation disturbance history was built for the study area from 1985 to 2013; however, the vegetation disturbances mapped were not a strong predictor of sage-grouse seasonal habitat-use. To evaluate effectiveness of conifer reduction treatments I used fecal pellet and in concert with radio-telemetry data. Increased sage-grouse use of conifer treatments was positively associated with sage-grouse presence in adjacent habitats (P = 0.018), percent shrub cover (P = 0.039), and mesic environments within 1000 m of treatments (P = 0.048). Sage-grouse use of conifer treatments was negatively associated with conifer canopy cover (P = 0.048) within 1000 m of treatments. To investigate sample bias related to individual bird behavior or capture trauma I monitored 204 radio-marked sage-grouse within the West Box Elder and Rich-Morgan- Summit SGMAs in Utah between January 2012 and March 2013. Sage-grouse that flushed one or more times prior to capture had higher brood (P = 0.014) and annual survival (P = 0.027) than those that did not. Sage-grouse that experienced more capture trauma had decreased annual survival probabilities (P = 0.04).