White-Nose Syndrome (WNS; Pseudogymnoascus destructans) is responsible for the regional population collapse of many cave-hibernating bat species, including the northern long-eared bat (Myotis septentrionalis), in eastern United States and Canada. I evaluated roosting behavior, habitat selection, and landscape-scale distribution of roosts of the northern long-eared bat during spring emergence and the early maternity season in Mammoth Cave National Park, Kentucky, USA, from 2015 to 2016. Logistic regression analysis comparing habitat features of roosts with random plots indicated selection of roosts reflected the costs of energetic demands by sex and reproductive status. Relative abundance of local bat species was assessed pre- and post-arrival of WNS in the Park during the summer season, with capture rates observed during the progression of WNS indicating that the fungal disease led to declines in the overall abundance of several bat species on the summer landscape, especially the northern long-eared bat. Distributional trends were quantified using spatial point pattern analysis which indicated that bats had clear roosting patterns associated with landscape level features and habitat resources. Monitoring bat populations regionally and at local scales will be imperative to helping conservation efforts for several bat species most affected by WNS.

Northern long-eared bat (Myotis septentrionalis) populations have declined sharply in recent years due to the introduction and rapid spread of White-nose Syndrome. This has prompted an urgent need for a greater understanding of their natural history in order to support the conservation of extant populations, particularly where forested day-roost and foraging habitats are being fragmented by development. Prior to 2006, with other Myotis species on the endangered species list, northern long-eared bats were understudied. In recent years, with the pressing concern to document the ecology of all cave bats affected by White-nose Syndrome on the landscape, researchers have now prioritized northern long-eared bat habitat needs, day-roost use, social dynamics, and barriers to gene flow. A relatively new discovery has been the numerous coastal populations that occur in smaller forest patches often surrounded by anthropogenic development. The goal of my research is to aid in informing future monitoring and management protocols that are specific to northern long-eared bats, particularly as they may shift from Threatened to Endangered under the Endangered Species Act. The chapters of this dissertation explore (1) similarities and differences between coastal and more documented upland habitat associations for populations of northern long-eared bats throughout the summer months through the use of acoustic detectors; (2) relationships between acoustic activity patterns and weather on an hourly basis for several bat species on Long Island, New York; and (3–4) day-roost characteristics and social network dynamics of a coastal northern long-eared bat maternity colony on Long Island, New York. I found that northern long-eared bat colonies have larger spatial footprints in – and a moderate tolerance for – forests in anthropogenically-developed areas than previously believed, though many natural history aspects of the bat were similar among coastal and interior populations. Installation of artificial day-roost structures such as bat boxes would likely be used and highly beneficial to colonies in these conditions, particularly to offset the deleterious effects of stochastic disturbance events on day-roost availability and to support social cohesion (and, by extension, reproductive success) for coastal maternity colonies. Additionally, I found that acoustic activity increased in riparian areas, or at sites closer to water features, and as proportion of forest coverage increased at a broad scale. However, immediately around the detector this relationship was reversed. Taken together, this demonstrates that riparian corridors, water features, and forest structural heterogeneity should explicitly be included in management guidelines. / Doctor of Philosophy / White-nose Syndrome is a fungal disease that has killed millions of cave bats in North America, including the northern long-eared bat (Myotis septentrionalis) that during the spring through fall uses forests for roosting and foraging habitat. As the disease continues to spread, and as forests are increasingly being lost to urbanization, it is imperative for researchers and land managers to understand the landscape needs of this species so that they can retain those forests that are used by this species. In addition to broader landscape needs, it is important to understand what specific forest stand to tree features the bats are using, how their social dynamics support their reproductive success, and how to best support habitat conditions foster mating in the fall swarm to avoid inbreeding. In recent years, more northern long-eared bats have been documented in coastal landscapes, suggesting that these areas might be more important to the conservation of the species than previously believed. In this dissertation, my aim was to explore how or why the landscapes in these coastal areas were being used by these bats both during nightly foraging and during the day as resting areas (day-roosts). Additionally, I explored how multiple species of bats were responding to weather conditions on the coast across seasons. Overall, I found that complex forests continue to be an important resource for northern long-eared bats, as well as water features whereby the latter should be more explicitly included in conservation management plans. Northern long-eared bats also show an ability to successfully use small forest patches within a larger urbanized landscape, including using human structures for day-roosts. Human structures might provide reliable, multi-year day-roosts in areas where extreme coastal weather events (hurricanes) occur or increase or as natural processes reduce the suitability of these forests to provide day-roosts. These structures, particularly artificial roost boxes could also support the social network of reproductive northern long-eared bats and thus maintain population stability to some extent.

Day-roost use by northern long-eared bat (Myotis septentrionalis) maternity colonies on the Fort Knox military reservation, Kentucky, USA, resulted in formation of non-random networks of roosts that exhibited a trend toward centralization. Centralization of day-roost networks was reflected in the social structure of colonies, which were characterized by dense associations among individuals within colonies. Social structure varied among colonies and appeared to be related to reproductive condition; colonies exhibited greater cohesiveness during parturition and lactation. Northern long-eared bat maternity colonies appeared to be exclusive, occupying distinct roosting areas with one or several areas receiving intense use. Day-roost removal simulations suggested a linear relationship between colony fragmentation and roost loss, and that loss of >20% of roosts is required to initiate colony fragmentation. Experimental hierarchical removal of day-roosts yielded results consistent with simulations, as removal of the single most-central (primary) roost had no impact on colony fragmentation, whereas removal of 24% of less-central (secondary) roosts resulted in partial network fragmentation. Patterns of colony day-roost and space use were similar pre- and post-removal treatments. Day-roost removal did not alter the number of roosts used by individual bats, but distances moved between roosts were greater in the secondary roost-removal treatment group. Day-roost characteristics largely were consistent pre-post treatment for both treatment groups. Historical data from an Indiana bat (Myotis sodalis) maternity colony revealed that this species also exhibits a non-random social assorting dynamic. Non-random social assortment resulted in a closely connected centralized network of day-roosts. Individuals within the Indiana bat maternity colony exhibited close social connections with colony members, but subgroups likely existed within the colony. Indiana bat day-roosting ecology appears flexible, as patterns of roost and space use differed substantially between years. Development of specific, but tactical, management approaches for individual colonies of both northern long-eared and Indiana bats may be possible. Such approaches would allow land managers to manage for both northern long-eared bat habitat and other objectives. However, the nature of targeted management approaches employed likely will depend on the unique forest context and dynamic within which individual colonies reside. / Ph. D.

Bat populations across North America are either facing new threats from white-nose syndrome (WNS) and wind energy development or have already experienced precipitous declines. Accordingly, researchers and managers need to know how to best monitor bats to document population and distribution changes, as well as where to look for persisting populations. Landscape-scale WNS impacts to summer bat populations are not well understood, and although acoustic monitoring is commonly used to monitor these populations, there is limited information about differences among acoustic detectors and the implications to managers thereof. My objectives were to model the relationship between WNS impact, influence of available hibernacula, and environmental factors for summer nightly presence of three WNS-affected bats and to compare how multiple models of acoustic detectors perform in terms of detection probability and nightly recorded bat activity. I collected acoustic data from 10 study areas across Virginia, West Virginia, Ohio and Kentucky to describe changes in nightly presence of WNS-affected bat species during summer 2017. During the same period of time, I compared five types of acoustic detectors at Fort Knox, Kentucky. My results show the potential efficacy of using a WNS impact-year metric to predict summer bat presence, and highlight which environmental variables are relevant for large-scale acoustic monitoring. Additionally, my findings suggest that each of the detector types tested would suffice for most research and monitoring activities, but standardization of detector type within the scope of a project or study should be encouraged. / Master of Science / Bat populations across North America are either facing new threats from white-nose syndrome (WNS) and wind energy development or have already experienced devastating declines. Accordingly, wildlife biologists need to know how to best monitor bats to document population and distribution changes, as well as where to look for remaining populations. Landscape-scale WNS impacts to summer bat populations are not well understood, and although acoustic technology is commonly used to monitor these populations, there is limited information about differences among acoustic detectors and the implications to managers thereof. My objectives were to model the relationship between WNS impact, influence of available bat hibernation caves, and environmental factors for summer nightly presence of three WNS-affected bats and to compare how multiple models of acoustic detectors perform in terms of detection probability and nightly recorded bat activity. I collected acoustic data from 10 study areas across Virginia, West Virginia, Ohio and Kentucky to describe changes in nightly presence of WNS-affected bat species during summer 2017. During the same period of time, I compared five types of acoustic detectors at Fort Knox, Kentucky. My results show potential viability of a WNS impact-year metric to predict summer bat presence, and highlight which environmental variables are relevant for large-scale acoustic monitoring. Additionally, my findings suggest that each of the detector types tested would suit most research and monitoring activities, but standardization of detector type within the scope of a project or study should be encouraged.

With declines in abundance and changing distribution of White-nose Syndrome (WNS)-affected bat species, increased reliance on acoustic monitoring is now the new "normal". As such, the ability to accurately identify individual bat species with acoustic identification programs has become increasingly important. Additionally, how bat distribution and habitat associations have changed at the local to sub-landscape scale in the post WNS environment is important to understand. The significance of these changes, relative to bat activity, may be based on the species-specific susceptibility to WNS. We used data collected from Fort Drum Military Installation, New York from the summers of 2003-2017 to analyze the accuracy of acoustic software programs, and assess the changes in relative bat activity, occupancy, and distribution induced by WNS. Our results indicate that continued acoustic monitoring of bat species, such as the little brown bat (Myotis lucifugus) in the Northeast, to assess ongoing temporal and spatial changes, habitat associations, and as a guide to direct future mist-netting should rely more on relative activity as the metric of choice. Furthermore, the continuous spread of WNS across North America will have strong negative effects on bat populations and communities, this study points to how individual species (both impacted and non-impacted) will respond to WNS. We believe that our results can help users choose automated software and MLE thresholds more appropriate for their needs to accurately address potential changes in communities of bat species due to impacts of WNS or other factors. / MS

Bats occupy diverse and unique niches and are regarded as important components in maintaining ecosystem health. They are major consumers of nocturnal insects, serve as pollinators, seed disperser, and provide important economic benefits as consumers of agricultural and forest pest insects. Bats have been proposed as good indicators of the integrity of natural communities because they integrate a number of resource attributes and may show population declines quickly if a resource attribute is missing. Establishing community- and population-level data, and understanding species interactions is especially important in changing landscapes and for species whose populations levels are threatened by outside factors of anthropomorphic disturbance from hibernacular visitation to energy production and fungal pathogens. For these reasons I have set out to establish habitat use patterns, detection probabilities, spatial and temporal occupancy, and investigate species interactions. This thesis is broken down into three distinct chapters each intended to be a stand-alone document. The first establishes the basic ecology from natural history accounts, provides an overview of the various sampling strategies, and gives a comprehensive description of the study area. The seconds sets out to identify the factors influencing detection probabilities and occupancy of six sympatric bats species and provide insight into habitat use patterns. The third examines spatial and temporal activity patterns and investigates species interactions. This study can provide understanding into the secretive and poorly understood patterns of free flying bats across the landscape. It can also deliver useful information to land managers regarding potential changes in landscape practices for the conservation of bat species. / Master of Science