This is a thesis about the spatial distribution of willow warbler, nightingale, blackbird, robin and grey flycatcher in Svealand and Götaland, that is the southern third of Sweden. It explores the possibilities of using statistics to describe the distribution and variation of birds in a given region.The data was collected by observation of birds on sites called standard routes, with 25 kilometres between them. The standard routes are the points in a grid net placed upon the map of Sweden. The purpose of standard routes is to represent the birds in Sweden both geographic and biotopological.The thesis compare the results from kriging, variogram and four alternative poisson regressions. In the end I come up with the information provided by kriging and variogram and which poisson regression that bests estimates the population sizes of the birds at a given site with information about year, mean temperature from January to May and what kind of environment or habitat the site consist of.

Master of Landscape Architecture / Department of Landscape Architecture/Regional and Community Planning / Melanie F. Klein / Greenways are an evolving landscape form, adapting to the needs of humans through time. Rooted in the work of Frederick Law Olmsted, greenways have progressed to become an international movement. Throughout this progression, three generations of greenways are recognizable—Generation 1: Parkways and Boulevards, Generation 2: Trail-Oriented Recreation, Generation 3: Multi-Objective (Searns 2002). Given the ideals of sustainability that drive our contemporary society, a fourth generation is poised to emerge. Responsive to sustainability, the fourth generation will be envisioned as a holistic system, structured within the frameworks of landscape ecology theory as developed by Richard Forman, and expanded with the progressive thinking of Jack Ahern and Kristina Hill. River North Greenway in Denver, Colorado is prime for transformation. Compromised by its industrial context, yet vitalized through the flow of the South Platte River and the infill development growing up the valley from Downtown Denver, River North will be re-defined as a holistic system. Utilizing the spatial pattern and process dynamics that define ecological theory (Forman 1996), abiotic, biotic, and cultural functions (Ahern 2007) will become the fundamental elements in strategizing the greenway as a multi-scalar dynamic mosaic (Hill 2005). River North Greenway will become a complexity of cultural activity and abiotic and biotic health, balancing programmed space with the enhancement and restoration of ecosystems. It will transform the City of Denver, enriching the connection between the city and its river, the people and nature. With its rich history and present potential, River North can become a timeless piece of Denver’s urban landscape, shaping meaningful human experiences and preserving nature within the built environment for future generations. In turn, it can propel the greenway movement towards a fourth generation.

Doctor of Philosophy / Department of Biology / Brett K. Sandercock / Greater prairie-chicken (Tympanuchus cupido) populations have been reduced by >70% since the turn of the 20th century due to large-scale conversion of native prairie habitats to cultivated agriculture and other human development. Although Kansas is considered a stronghold for greater prairie-chickens, statewide populations have declined >30% in the last 30 years. Goals of this dissertation were to determine the demographic mechanisms for apparent population declines and evaluate how regional variations in landscape composition and grassland management affect the demography, habitat use, life-history, and population viability of three populations of greater prairie-chickens. First, I found that, despite high reproductive potential, poor reproductive success prevented populations from being self-sustaining. All three populations were projected to decline but finite rates of population declines were different among populations (λ = 0.49, 0.54, and 0.74). I found that grassland fragmentation and rangeland management practices influence nearly every aspect of greater prairie-chicken population ecology and dynamics. A population in a contiguous prairie landscape managed with annual spring burning and intensive early stocking of cattle (South) was characterized by delayed breeding, low nest and brood survival (0.08–0.18 and 0.27, respectively), high annual survival of mature females (0.64–0.71), projected age-ratios heavily skewed toward adults, and longer generation times. Conversely, a population in grasslands heavily fragmented by cultivation and managed with longer fire-return intervals and moderate grazing (Smoky) initiated nests earlier, had higher nest and brood survival rates (0.16–0.31 and 0.34, respectively), produced significantly larger eggs, and had low annual survival (0.34–0.42) and shorter generation times. A site with intermediate levels of fragmentation, burning and grazing (North) had intermediate demography. Finite population change was more sensitive to changes in adult survival at all sites, but the relative influence of fecundity parameters on projected population change was not similar among study populations. Data indicate that differences in rates of decline among populations were largely due to variation in adult survival mediated by human landscape alteration. Human-mediated changes to grasslands impact the demography and viability of prairie-chicken populations, influence population sensitivities to changes in vital rates, and mediate changes in the life-history strategies of a grassland-sensitive species.

published_or_final_version / Geography / Doctoral / Doctor of Philosophy

This study describes and analyses cattle management in Tlokweng Sub District. Two methods were used. The two are households' interviews and a system dynamics STELLA model called the Rain Land Cattle model, which was adapted from the 1990 Braat and Opschoor model. Ninety households, 61% of the 1991 households in the study area, were interviewed. All the households had arable fields and fifty nine percent had cattle. The Rain Land Cattle model uses 52 parameters to predict several cattle management factors, which include rainfall, stocking rate, total grazing area and livestock water availability. The model explored the use of parameters to relate water availability to grazing area and show the seasonality of the water source. Sixty two percent of the household had access to an ideal livestock water source. Cattle graze from the 5000 hectares of arable area for four months after harvesting. This seasonal grazing, optimises the uses of the grazing resource in the small sub - district. The model simulated a S and 20 percent permanent grazing land loss. Such a grazing land loss, increased the stocking rate, decreased the carrying capacity and cast doubt on sustainable cattle production. The model shows that the stocking rate is chronically greater than the carrying capacity. Most households acknowledged that there was land pressure due to the loss of grazing land. A drier climate scenario will lead to a loss of seasonal grazing, reduced livestock water, which will increase cattle emigration and cause cattle management problems. The model is exploratory; it needs to be validated. It is easily understood, adaptable to other communal areas, and identifies the most influential factors in cattle management. The livestock water parameters functioned reliably in the model. Based on the understanding of the cattle management derived from this study, more fenced grazing land is unlikely to improve the cattle management in the area.

The western Amazon is known to be one of the most biologically diverse regions in the world, yet information about the spatial distribution of that biodiversity and the processes governing its distribution remains scarce. An improved understanding of those biogeographic patterns and processes can inform conservation and development planning in areas where anthropogenic landscape change is ongoing. Spatial components of biodiversity are known to be influenced by historical and present-day physical and human geographic processes. There is evidence that major Amazonian rivers form the boundaries of biological regions, at least for birds. Other factors that may influence bird species composition include the dispersal limitations of individual species, forest plant species composition and structure, topography, forest fragmentation, and hunting. Sites where bird species composition was measured in this study represented mature, upland forest on both sides of the Amazon River, and a range of non-flooded forest types, as indicated by soil and plant surveys. Bird species compositional variation was closely correlated with variation in plant species composition, human disturbance associated with forest fragmentation, and position north or south of the Amazon River. The strongest differences were between opposite sides of the river, even though local environments, including plant composition, were not different on the two sides. This strongly suggests that historical biogeographic factors, rather than present-day environmental gradients, are responsible for bioregional boundaries at Amazonian rivers. The difference between plant and bird distributions at this scale underscores the pressing need to re-evaluate general notions of bioregional complexity and pattern in the Amazon basin. Locally, the influence of habitat fragmentation on animal communities, including reduced species richness, was confirmed. The influence of local floristic variation is of particular importance due to its ubiquity across western Amazonia. Thus, understanding the distributions of soils and vegetation is critical for explaining Amazonian animal diversity. The use of these factors to model bird community heterogeneity contradicts assumptions that the processes shaping Amazonian animal community diversity are too complex to measure efficiently, and their use contributes a new understanding of the dimensions of that diversity. / text

Compared to natural forests, fast-growing plantations of exotic species such as Pinus radiata are often perceived as marginal habitat or unsuitable habitat for most native species. By studying Coleoptera (beetles) in a variety of landscape elements (pasture, native forest and different aged Pinus radiata stands) in a highly modified and fragmented landscape in New Zealand I aimed to determine the value of exotic plantation forests for native biodiversity, and how these species are affected by different sized clearfell harvest areas. Pitfall trap sampling of beetles showed that plantation forest stands can provide suitable complimentary habitat to native forest for many species. Rarefied species richness of Carabidae, Scarabaeidae and Scolytinae was not significantly different between habitats, however, habitat types differed significantly in their beetle community composition. Comparing different production habitats, Pinus radiata stands had a beetle community composition most similar to native forest. However, a small minority of species, e.g., Dichrochile maura, were restricted to native forest habitat highlighting the importance of retaining indigenous ecosystems within plantations. Unlike human modified habitats, native forests did not provide suitable habitat for exotic species. Clearfell harvesting is controversial and its impact on biodiversity is a key constraint for many forest certification programs, such as that administered by the Forest Stewardship Council (FSC). Despite this, no replicated manipulative experimental studies of the impact of different sized clearfell harvest areas on biodiversity have been undertaken at scales relevant to the New Zealand forest industry. One potential model of the impact of different clearfell harvest sizes is the concept of a threshold size. A threshold scenario may occur where clearfell harvest impacts increase at a rate disproportionate to the change in clearfell size over a small range of harvest areas, but impacts remain relatively unchanged either side of the threshold zone. I sampled Coleoptera in experimentally created 0.01, 0.05, 0.5, 5.0, 50 and 500 ha clearfells within Pinus radiata plantations in the central North Island of New Zealand. The wide range of clearfell harvest sizes, including some very small areas, such as 0.01 ha was instigated in an attempt to document potential clearfell harvest size thresholds. Rarefied native beetle species richness was higher in harvest areas compared to adjacent mature plantation stands. The beetle species richness in 5 ha and 500 ha harvest areas was significantly greater species than that in small 0.01 - 0.5 ha harvest areas. Although, the high beetle diversity recorded in 500 ha clearfells should be treated with caution due to confounding spatial autocorrelation. The degree of change in beetle community composition increased with increasing clearfell harvest area. Beetle assemblages in large harvest areas were less similar to their paired adjacent mature forest than smaller harvest areas. Although, constrained multivariate ordination techniques did show a short-term change in beetle species composition between recently clearfelled harvest areas of as little as 0.05 ha and adjacent mature P. radiata stands. The colonisation by open-habitat disturbance-adapted species was a key driver of this change, some species dispersed into clearfelled stands in significant densities within days post-harvest. Overall, there were no distinct short-term trends to the change in species richness as a function of increasing harvest area that would suggest an ecological impact threshold response. If short-term outcomes of clearfell harvesting are ameliorated by successful recolonisation, the long-term spatial arrangement of different aged stands becomes more important for the maintenance of biodiversity at the landscape level than short-term consequences of harvesting. By sampling selected beetle taxa in 1, 2, 4, 8, 16 and 26 year-old stands, I found that the abundance of seven out of eight of the species selected for analysis recovered to levels similar to those in adjacent mature forest within the timeframe of a single harvest rotation. Individual species utilised different aged stands, indicating different life-history strategies. For example, open-habitat, disturbance-adapted species such as Cicindela tuberculata and Sitona discoideus were prominent in young stands, and forest species such as Pycnomerus sophorae and Paracatops phyllobius were highly abundant in older stands. These alternative life-history strategies highlight the benefits of maintaining a mixture of different aged stands to increase biodiversity at the landscape level. This thesis fills an important gap in our knowledge of biodiversity in production landscapes. I show that plantation forests have value as complimentary habitat to native forest and they make an important contribution to the maintenance of biodiversity at the landscape level. Although clearfell harvesting is a severe disturbance to the forest ecosystem, the long-term recovery of beetle populations suggests that harvesting is not the key limiting factor to the enhancement of biodiversity in the plantation forests studied. This unusual situation is possibly the result of prior land-use history, as many plantations were established on degraded pastoral land, and harvest-sensitive species are unlikely to have survived this initial land-use change. As such, the severity of the long-term impacts of clearfell harvesting on biodiversity are likely to be context specific and will vary accordingly. The importance of spatial heterogeneity of habitat elements, including different aged plantation stands and native forest remnants, needs to be investigated in more detail to determine what limits biodiversity in this plantation landscape. Key points to consider are the proximity to, and proportion of, native forest cover in the landscape and the degree of connectivity among native remnants. It is these landscape-level attributes that may determine biodiversity at a regional scale, and more emphasis should be placed on landscape scale factors and there interaction with stand specific forest management practices. For example, the spatial mosaic of harvesting areas may need to be of a finer-scale when there are fewer native remnants within the landscape.

This thesis examines the ecology, parasites and pathogens of three insectivorous bat species in Wytham Woods, Oxfordshire; Myotis nattereri (Natterer’s bat), M. daubentonii (Daubenton’s bat) and Plecotus auritus (Brown long-eared bat). The population structure was assessed by monitoring associations between ringed individuals, utilising recent advances in social network analysis. Populations of both M. daubentonii and M. nattereri were found to subdivide into tight-knit social groups roosting within small areas of a continuous woodland (average minimum roost home range of 0.23km2 and 0.17km2 respectively). If this population structure is a general attribute of these species it may make them more sensitive to small scale habitat change than previously thought and has implications for how diseases may spread through the population. M. daubentonii had a strong preference for roosts close to water, away from woodland edge and in areas with an easterly aspect. The factors driving roost choice in M. nattereri and P. auritus remain elusive. The segregation of M. daubentonii into bachelor and nursery colonies was not a result of the exclusion of males from roosts close to water by females, or variation in microclimate preferences between the sexes, as was predicted. Body condition (weight/forearm length) was correlated with host characteristics including age and reproductive status, and weather variables. Astroviruses and Coronaviruses, which have characteristics typical of zoonotic viruses, were identified in UK bat species for the first time. Coronaviruses identified formed species-specific clades while Astroviruses were highly diverse. Though not closely related to human viruses these are potential zoonotic diseases of the future. Models of Coronavirus and ectoparasite distribution suggest individual attributes (e.g. sex and age) and population structure (e.g. the formation of nursery and bachelor colonies) are important predictors of parasite and pathogen prevalence. This study characterises a system that offers many opportunities for future research including studies of sociality, disease modelling and conservation management.

Urbanisation is a key driver in the loss, fragmentation and modification of natural habitats resulting in the global loss of biodiversity. As the human population, and consequently the rate of urbanisation, continues to increase exponentially it is important to understand how to sustain and enhance biodiversity within the built environment. Cities comprise a complex assortment of habitat types yet relatively little is known of how its composition and spatial configuration can influence species presence or foraging activities. It is therefore necessary to examine habitat use and biodiversity patterns at multiple spatial scales to fully understand how species are responding to the urban matrix. There are few other orders of animals that are as strongly associated with people as bats (Chiroptera); for some bat species human habitations provide roosts and adaptations of the environment provide food sources. However bat species richness generally declines with increasing urbanisation indicating that many species are not able to persist in highly urbanised areas. In this thesis, I show that the behaviour, habitat preferences, and distribution of bats are strongly influenced by the built environment at both a local and landscape scale. Although many animal species are known to exhibit sex differences in habitat use, adaptability to the urban landscape is commonly examined at the species level without consideration of potential intraspecific differences. I found that female Pipistrellus pygmaeus show greater selectivity in foraging locations within urban woodland in comparison to males at both a local and landscape scale. There was a lower probability of finding females within woodlands which were poorly connected, highly cluttered, with a high edge: interior ratio and fewer mature trees. The results have important implications for our understanding of how to manage areas for breeding females and highlight the need to supplement acoustic monitoring with trapping data to assess sex differences in habitat use. Determining how morphological or behavioural traits can influence species adaptability to the built environment may enable us to improve the effectiveness of conservation efforts. The morphological similarities between P. pygmaeus and P. pipistrellus suggest that both species should respond similarly to the urban matrix, however I found differential habitat use occurring within a variety of urban habitats (e.g. woodland and waterways) and at a landscape scale. In urban woodland there was a higher probability of P. pygmaeus activity relative to P. pipistrellus in woodlands with low clutter and understory cover which were surrounded by low levels of built environment. Many bat species are strongly associated with aquatic or adjacent riparian habitats yet we know little about the utilisation of urban waterways by bats. After surveying urban waterways throughout the UK, I was able to show that the built environment can negatively affect a variety of bat species from the riparian zone up to 3km from a waterway. This indicates that beneficial urban waterway rehabilitation schemes for bats require management at multiple spatial scales, from retaining a vegetated riparian zone at the local scale to highlighting the necessity for conservation funding to be spent on the implementation of landscape scale environmental improvement schemes that encompass the entire urban matrix. Undertaking surveys to confirm species presence or to estimate population sizes can be difficult, particularly for elusive species such as bats. I was able to demonstrate a variety of ways to increase surveying efficiency (e.g. the use of an acoustic lure to increase bat-capture rate) and a significant relationship between bat activity and the relative abundance of certain species of bat which can maximise the knowledge of diversity in an area whilst minimising wildlife disturbances. Urbanisation has also had strong negative effects on many insect groups, such as moths, which are important components of the diets of many bat species. I found that woodland vegetation characteristics were more important than the surrounding landscapes in determining the abundance, species richness, and species diversity of moth assemblages within urban woodland. This indicates that management at a local scale to ensure provision of good quality habitat may be more beneficial for moth populations than improving habitat connectivity across the urban matrix. The findings presented in this thesis have important implications for our understanding of the adaptability of species to the built environment and for the management and monitoring of bat populations. It also highlights that even common bat species are negatively affected by urbanisation and much greater attention should be paid to securing their future within the urban landscape.

The environmental problems facing the world today are largely attributable to anthropogenic activities and landscape change. Addressing these challenges in an evidence-based way requires an understanding of precisely how species and ecosystems are responding to human impacts. Discerning linkages between stressors and their ecological repercussions, and using this to inform conservation, can be challenging due to the complexity and uncertainty of ecological research. I focused on the responses of five wide-ranging large mammal species – gray wolf (Canis lupus), Canada lynx (Lynx canadensis), coyote (Canis latrans), white-tailed deer (Odocoileus virginianus) and moose (Alces alces) – to human footprint (measure of human infrastructure and landscape change), using 12 years (2001-2013) of snowtrack surveys conducted across the boreal forest of Alberta. I explored three key challenges to discerning the linkages between ecological dynamics and management actions. First, I asked whether the direction and magnitude of species responses vary depending on the spatial extent and grain of the study. Second, I asked whether these species respond more strongly to individual footprint features or to the cumulative effects of footprint (measured as total footprint), and whether responses to footprint are consistent across species. Third, I evaluated the utility of thresholds for large mammal management and asked whether there is evidence for consistent threshold responses to total footprint across scales. In addressing the first two questions, I evaluated a set of generalized linear mixed effects models (GLMM) relating the relative abundance of each species to individual and cumulative effects of human footprint, using an information-theoretic approach. I compared the direction of species responses across our regional study area (approximately 400,000 km2) to those reported in previous smaller-extent studies (median 1,525 km2), and compared responses across three spatial grains (250m, 1500m, and 5000m transect buffers). In addressing the third question, I conducted a review on the utility of ecological thresholds, described as abrupt changes in the response to a continuous driver, for large mammal management. I further tested for thresholds in species responses to total footprint by comparing linear models (logistic regression) to piecewise regression models. I compared threshold values between two grains (approximately 33km2 - 1500m transect buffer, and 5500km2 - grouping transects into clusters), and across four regions (boreal forest extent, three landscape planning units). I found that the direction of species responses varied with spatial extent, but not grain, and that species responded strongly to a broad suite of footprint features, indicating the need to manage for cumulative effects. Despite the appeal of ecological thresholds, using these as targets is challenging and the success of doing so has rarely been evaluated. I found threshold models to be better supported than linear ones across species, but due to variability and uncertainty in threshold values, the results are more suited as guidelines or hypotheses to be further tested, as opposed to specific management targets. Translating research on complex ecological systems into management actions is a continuing challenge, yet, ongoing biodiversity monitoring and adaptive management may refine our existing tools, and ultimately lead to better environmental stewardship. / Graduate / 2017-09-05 / 0329