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In this study, two hydrologic models were applied to the mountainous Back Creek catchment, located in the headwaters of the Roanoke River in Southwest Virginia. The two models employed were HEC-1, an event based lumped model, and TOPMODEL, a continuous semi-distributed model. These models were used to investigate (a) the issue of spatial scale in hydrologic modeling, and (b) two approaches to modeling, continuous versus event based. Two HEC-1 models were developed with a different number of subareas in each. The hydrographs generated by each HEC-1 model for a number of large rainfall events were analyzed visually and statistically. No observable improvement resulted from increasing the number of subareas in the HEC-1 models from 20 to 81. TOPMODEL was applied to the same watershed using a series of different size grid cells. The first step in applying TOPMODEL to a watershed involves GIS analysis which results in a raster grid of elevations used for the calculation of the topographic index, ln(a/tan b). The hydrographs generated by TOPMODEL with each grid cell size were compared in order to assess the sensitivity of TOPMODEL hydrographs to grid cell size. An increase in grid cell size from 15 to 120 meters resulted in increased values of the watershed mean of the topographic index. However, hydrographs generated by TOPMODEL were completely unaffected by this increase in the topographic index. Analyses were also performed to determine the sensitivity of TOPMODEL hydrographs to several model parameters. It was determined that the parameters that had the greatest effect on hydrographs generated by TOPMODEL were the m and ln(To) parameters. The modeling performances of the event based HEC-1 and the continuous TOPMODEL were analyzed and compared visually and statistically for a number of large storms. The limited number of storms used to compare HEC-1 and TOPMODEL makes it difficult to determine definitively which model simulates large storms better. It does appear that perhaps HEC-1 is slightly superior in that regard. TOPMODEL was also executed as an event based model for two single events and the resulting hydrographs were compared to the HEC-1 and continuous TOPMODEL results. Both HEC-1 and TOPMODEL (when used as a continuous model) simulate large storms better than TOPMODEL (when used as an event based model). / Master of Science

Despite their difference in body size and morphology, the moose (Alces alces) andthe mountain hare (Lepus timidus) sustain themselves during winter on similar plantspecies and plant parts in in subarctic environments, namely apical twigs ofmountain birch (Betula pubescens ssp. czerepanovii). Herbivores must select areas anditems of food that provide sufficient intake rates and food nutritional quality whilebalancing this against their intake of dietary fiber and potentially detrimental plantsecondary metabolites. This selection takes place simultaneously at multiple spatialscales, from individual plants and plant parts to patches of food and parts of the wider landscape. While the herbivores must consider their need for food to sustaindaily activities, for body growth and reproduction it is also necessary to avoid predators and harsh environmental conditions. For managers, an understanding of key factors for animal foraging distributions is pivotal to reach intended goals ofmanagement and conservation plans. Knowledge in this area is also important formodels to make accurate predictions of foraging responses of herbivores to resource distributions. The mountain birch forest displays a naturally heterogeneous distribution of trees and shrubs which presents herbivores with a challenge to findgood feeding areas. In an investigation of the spatial distribution of moose browsing on birch and willows (Salix spp.) in two winter seasons separated in time by 14 years,it was found that moose browsing patterns in 1996 were correlated to those observed in 2010. It was also found that moose browsing was spatially clustered within the same distances (1000-2500 m) as densities of willow and birch, but at other spatial scales, browsing was mostly randomly distributed. It was concluded that foragedensity is a cue for moose but only at certain spatial scales. Similarly, a comparison of foraging distribution by hare and moose showed that high birch density was a key factor for both species. In spite of this, hares and moose used different parts ofthe same environment because they respond to food resource distribution at different spatial scales. Hares fed from smaller plants, and focused their foraging activity on smaller spatial scales than moose. These results emphasize the importance of taking into account the distribution of food resources at spatial scales relevant for each species in plans for conservation and management. In an experimental study it was found that intensified browsing on natural forage by mountain hares can be induced locally through placement of food. The induced browsing varied with the amount and quality of the added food, but also with thedensity of natural food plants and natural foraging distribution by hares. Finally, ina last experiment habitat preference of mountain hares across edges between open and forested areas was studied. The results were not consistent; hares utilized baitto a greater extent within forested areas than bait placed on a nearby lake ice, butbait on mires and heaths was either preferred over bait in nearby forest, or utilizedto a similar extent. A possible explanation is that hares have knowledge of their environment such that both forested areas and subarctic mires and heaths are partof its natural home range, whilst the extreme environment on the lake ice is not. During recent decades arctic areas have had an increase in vegetation density andwill be affected by future climate warming and therefore, factors that determineforaging ecology of key herbivores need to be identified. This thesis sheds some light on these factors in relation to spatial scale and forage distribution for two high profile herbivores in the subarctic.

Studies have found that the level of association between an area-level covariate and an outcome can vary depending on the spatial scale (SS) of a particular covariate. However, covariates used in regression models are customarily modeled at the same spatial unit. In this dissertation, we developed four SS model selection algorithms that select the best spatial scale for each area-level covariate. The SS forward stepwise, SS incremental forward stagewise, SS least angle regression (LARS), and SS lasso algorithms allow for the selection of different area-level covariates at different spatial scales, while constraining each covariate to enter at most one spatial scale. We applied our methods to two real applications with area-level covariates available at multiple scales to model variation in the following outcomes: 1) nitrate concentrations in private wells in Iowa and 2) body mass index z-scores of pediatric patients of the Virginia Commonwealth University Medical Center. In both applications, our SS algorithms selected covariates at different spatial scales, producing a better goodness of fit in comparison to traditional models, where all area-level covariates were modeled at the same scale. We evaluated our methods using simulation studies to examine the performance of the SS algorithms and found that the SS algorithms generally outperformed the conventional modeling approaches. These findings underscore the importance of considering spatial scale when performing model selection.

Understanding the factors affecting species distribution and at what scale a species respond to these factors is a major challenge in conservation biology. I studied the distribution and abundance of a monophagous shield bug Canthophorus impressus at three spatial scales: host plants, patches and circular landscapes (area ca. 3 km2), to determine the relative importance of the different spatial scales and how habitat quality, amount and spatial configuration affect the distribution of C. impressus. Influence of habitat characteristics on occurrence and abundance of the bug was analysed with separate generalised linear mixed models. The data show that effects of habitat quality and amount dominate over configuration in determining the distribution of C. impressus. The bug prefers large host plants in warm conditions on a plant scale and abundant host plants in a landscape scale, whereas patch scale was not important for the distribution of the bug. Management should aim to preserve sites with abundant host plants and promote large host plants with low to moderate grazing. To secure sufficient number of host plants in the landscape, high densities of suitable habitat such as semi-natural grasslands should be preserved. This study suggests that analyses of multiple spatial scales are crucial for identifying appropriate actions for successful conservation of species living in fragmented habitats.

In Soil and Water Assessment Tool (SWAT), oftentimes, Critical Source Area (CSA), the minimum upstream drainage area that is required to initiate a stream, is used to subdivide a watershed. In the current literature, CSA has been used as a trial and error process to define the subwatershed levels. On the other hand, the ongoing collaboration of the United States Environmental Protection Agency Office of Water and the United States Geological Survey has promoted a national level predefined catchments and flowlines called National Hydrography Dataset (NHD) Plus to ease watershed modeling in the United States. The introduction of NHDPlus can eliminate the uncertain nature in defining the number of subwatersheds required to model the hydrologic system. This study demonstrates an integrated modeling environment with SWAT and NHDPlus spatial datasets. A spatial tool that was developed in a Geographical Information System (GIS) environment to by-pass the default watershed delineation in ArcSWAT, the GIS interface to SWAT, with the introduction of NHDPlus catchments and flowlines, was used in this study. This study investigates the effect of the spatial size (catchment area) of the NHDPlus and the input data resolution (cell/pixel size) within NHDPlus catchments on SWAT streamflow and sediment prediction. In addition, an entropy based watershed subdivision scheme is presented by using the landuse and soil spatial datasets with the conventional CSA approach to investigate if one of the CSAs can be considered to produce the best SWAT prediction on streamflow. Two watersheds (Kings Creek, Texas and Sugar Creek, Indiana) were used in this study. The study shows that there exists a subwatershed map that does not belong to one of the subwatershed maps produced through conventional CSA approach, to produce a better result on uncalibrated monthly SWAT streamflow prediction. Beyond the critical threshold, the CSA threshold which gives the best uncalibrated monthly streamflow prediction among a given set of CSAs, the SWAT performance can be improved further by subdividing some of the subwatersheds at this critical threshold. The study also shows that the input data resolution (within each NHDPlus catchments) does not have an influence on SWAT streamflow prediction for the selected watersheds. However, there is a change on streamflow prediction as the area of the NHDPlus catchment changes. Beyond a certain catchment size (8-9% of the watershed area), as the input data resolution becomes finer, the total sediment increases whereas the sediment prediction in high flow regime decreases. As the NHDPlus catchment size changes, the stream power has an influence on total sediment prediction. However, as the input data resolution changes, but keeping the NHDPlus catchment size constant, the Modified Universal Soil Loss Equation topographic factor has an influence on total sediment prediction.

Rivers are hierarchical networks that integrate both large and small scale processes within catchments. They are highly influenced by variation in flow and are characterised by strong longitudinal movement of materials. I conducted an extensive literature review that indicated braided rivers lie at the upper end of the river complexity gradient due to the addition of strong lateral and vertical connectivity with their floodplains. The management of these rivers requires an understanding of the connective linkages that drive complexity, however in developed regions few braided river systems remain intact. The large number of relatively pristine braided rivers in New Zealand provided a unique opportunity to study physical and biotic patterns in these large dynamic systems. Initially I examined horizontal connectivity through patterns in regional and local diversity in eleven braided rivers in the North and South islands of New Zealand. Subsequently, the next component of my thesis focused on vertical connectivity through intensive investigations of energy pathways and the recipient spring stream food-webs. The eleven river survey included sampling of multiple reaches and habitats (main channels, side braids, spring sources, spring streams and ponds) and confirmed the importance of lateral habitats to invertebrate diversity. However, I found that large spatial scales made a greater contribution to diversity than small scales, such that major differences occurred between rivers rather than habitats. This result suggested either a role for catchment-scale factors, such as flow, or biogeographic patterning. Subsequent analysis of the relationships between invertebrate diversity and the physical environment indicated strong regulation by flow variability, but also biogeographic community patterns. Braided rivers are clearly disturbance dominated ecosystems, however the effects of disturbance are manifest in different ways across the riverscape. The role of vertical hydrological connectivity in linking the different components of the floodplain was investigated by tracing carbon pathways from the terrestrial floodplain to spring-fed streams and their communities. Using δ13C isotope signatures it was possible to show that inorganic carbon in groundwater was derived from terrestrial vegetation and subsequently incorporated into spring stream food-webs. However, the degree to which a stream community uses groundwater as opposed to allochthonous carbon is affected by the successional stage of riparian vegetation, a function of the shifting habitat mosaic that is regulated primarily by flow variation and sediment dynamics. In summary, the structure of braided river ecosystems is regulated primarily at the catchment scale, but connectivity at smaller scales plays an important role in determining ecological structure and function.

This dissertation examines the relationship between abundances of Roanoke darter, Roanoke logperch, and black jump rock and availability of stream habitat features at three spatial scales in two reaches of the Roanoke River, Virginia. The utility of underwater observation as a method of estimating benthic fish densities is also assessed. Distributions of perpendicular sighting distances indicate models assuming equal sighting probability are not appropriate for benthic species but distance sampling models assuming decreased sighting probability with increased distance from observers provide reasonable alternatives. Abundances estimated using two distance sampling models, a strip transect model, and a backpack electroshocker were strongly correlated. At the microhabitat scale (45 m² cells), differential use of depth, velocity, substrate, and siltation level by all three species during summer low flows was evident. Habitat use characteristics were not transferable, as depths and velocities associated with high fish densities varied between reaches. Univariate and multivariate habitat suitability indices gave similar rankings for combinations of the four habitat variables, but site suitabilities based on these indices were poor predictors of fish abundance. Habitat cells were not selected independently of surrounding habitat characteristics, as fish densities were highest in target cells adjacent to cells with preferred microhabitat characteristics. Roanoke darter and black jumprock abundances were highest at sites where preferred microhabitat patches were non-contiguous while contiguity had no effect on logperch abundance. Multiple regressions showed area of suitable habitat and patch contiguity accounted for 42 %, 34 %, and 33 % of variation in darter, logperch, and jumprock abundances, respectively. Estimates of area of target riffles, area of pools and riffles upstream and downstream of target riffles, and depth, velocity, and substrate characteristics of pools and riffles immediately upstream and downstream of target riffles were obtained. Fish densities were correlated with at least one measure of proximal habitat for all three species. Significant multiple regression models relating fish density to adjacent habitat unit characteristics were also obtained, but the explanatory power of adjacent unit variables varied among small, medium and large riffles and among species. In summary, fish abundance was related to habitat at all spatial scales but explanatory power was limited. / Ph. D.

Habitat loss and fragmentation drives the current extinction crisis. The processes through which it affects biodiversity, however, are complex and poorly understood. This is especially true for spatially complex regions that comprise a mosaic of land-use types, which often range from protected areas to dense human settlements. In such human-modified landscapes, it is important to determine the extent and impact of changing land-use patterns on biodiversity if we are to meet conservation targets or regain ecosystem services. My analyses of coastal forests in KwaZulu-Natal suggest that extensive loss of forests (82%) incurred an extinction debt, modelled to match the 11 bird species now listed as threatened locally. Forest fragments are now also smaller, fewer, further apart and more encroached by human land uses than in the past. Yet, species interactions with the gradient of habitat conditions that now surround forest fragments may have forestalled the realisation of predicted extinctions. I found that natural matrix habitats adjacent forest fragments (e.g. grasslands and woodlands) may facilitate dispersal, enable species spillover from forest fragments, and buffer forest interiors from changes in abiotic conditions associated with high contrast matrix habitats (e.g. agricultural plantations). However, when natural matrix habitats were transformed, these processes were disrupted, which suggest that the effect of landscape change on coastal forest diversity may stretch beyond forest loss per se and the deterministic extinctions predicted by conventional species-area relationships. Next, I determined that the response of different bird species to habitat fragmentation parameters (i.e. area, connectivity and matrix habitats), depended on life-history traits such as body size, feeding guild and habitat specialization. Extinction risk was, however, not a function of species traits or the fragmentation parameter species responded to. This means that a conservation approach that only focuses on restoring a single fragmentation parameter (e.g. area) may not be successful in halting predicted extinctions, simply because multiple factors may determine extinction risk in coastal forests. The interpretation of biodiversity patterns in fragmented landscapes may, however, also be influenced by spatial scale. I therefore used a fractal-based sampling design to test how sampling at fine, intermediate and coarse scales influences (1) beta diversity of and (2) inferences from the modelled contribution of niche- versus dispersal-based assembly processes in structuring tree and bird assemblages. I showed that inferences from beta diversity are scale dependent. As a result, studies with similar sampling effort and temporal sampling protocol, but with different sampling grains are likely to report dissimilar ecological patterns, which may ultimately lead to inappropriate conservation strategies. This thesis provides information of how land-use changes impact on biodiversity patterns and derived processes in a human-modified landscape. It also highlights some conservation opportunities in the coastal forest landscape mosaic, where conservation and restoration actions should focus on both forest fragments and on the surrounding matrices. The conservation of natural matrices may buffer forest communities from impacts associated with high contrast habitat edges, enhance natural plant regeneration through species spillover, provide important linkages between forest fragments, boost regional diversity and allow coastal forests to track environmental change under changing climatic conditions. / Thesis (PhD)--University of Pretoria, 2014. / tm2015 / Zoology and Entomology / PhD / Unrestricted

At the Fort Bragg Military Installation, an army base in North Carolina, the habitat associations for the breeding bird species and the effects of the current prescribed fire program on the avifauna are virtually unknown. Fort Bragg encompasses one of the largest, fire-dependent longleaf pine systems existing today and is a mosaic of forested habitats. I used bird count data collected during 1994-1997 at 50-m fixed radius point count stations to examine bird species-habitat relationships in relation to fire treatment (i.e., fire intense longleaf pine woodlands versus fire suppressed mixed pine-hardwood and hardwood forests) and a riparian-upland habitat gradient, and at multiple spatial scales (i.e., the microhabitat and landscape). I used two-way factorial analyses to test for the effects of fire treatment and the riparian-upland habitat gradient on total bird abundance, species richness, and species relative abundance. To examine species-habitat associations at multiple spatial scales, I measured vegetation characteristics at a 50-m radius microhabitat scale, and I quantified landscape structural attributes at a 300-m to 1500-m radius landscape scale using a GIS database and the spatial analysis program FRAGSTATS. I then used logistic regression to determine which microhabitat and landscape variables were associated with the probability of occurrence for each species and which spatial scale was of greater relative importance to a species' occurrence. Finally, I tested logistic regression (LR) models and multiple linear regression (MLR) models, specific to the microhabitat scale, with independent data to evaluate their usefulness at predicting the occurrence and relative abundance for several breeding bird species. Total bird abundance did not vary across fire treatment and species richness may be only slightly greater in fire suppressed habitats, even though this habitat offered greater structural complexity than the park-like longleaf pine, fire intense habitats. Both total bird abundance and species richness were highest within the riparian habitat of streamhead pocosins, which offered distinctive vegetative characteristics otherwise lacking in this landscape. The fire treatment and riparian-upland habitat gradient also were greatly associated with the relative abundance of many species. Four bird species assemblages were defined based on the relative abundance patterns across fire treatments and the riparian-upland gradient: longleaf pine, fire suppressed, drain (i.e., riparian habitat), and generalist assemblages. Continued longleaf pine restoration using growing season prescribed fire likely will cause a decline in species of the fire suppressed assemblage in mixed pine-hardwood and hardwood forests, including many Neotropical migrant songbirds, but will greatly benefit members of the longleaf pine assemblage, such as the Red-cockaded Woodpecker, Brown-headed Nuthatch, Prairie Warbler, and Bachman's Sparrow. Breeding bird distributions in this fire-influenced, forest-dominated system were associated with attributes at both microhabitat and landscape spatial scales, though microhabitat attributes generally were of greater importance for the occurrence of most species. Microhabitat variation associated with the fire management gradient (intensely burned habitat versus fire suppressed habitat) and the riparian-upland gradient were the most frequent predictors in the species-habitat models. These results are similar to other studies documenting that microhabitat features were more influential than landscape features for birds in a naturally patchy or forest-dominated landscape. The microhabitat LR (probability of occurrence) models performed best in presence/absence classification when tested with the same data used for model development (cross-validation tests), and the LR and MLR (relative abundance) models performed better for an independent two-year data set compared to an independent one-year data set (validation tests). Although most MLR models were not significantly biased when tested with an independent two-year data set, these models had relatively low precision, suggesting they can be used to predict species relative abundance across a large area but they may not be sensitive to changes in abundance at individual count stations. These model validation results suggest that modeling species occurrence, rather than both occurrence and relative abundance, would have been sufficient to describe general species-habitat associations and to produce reliable, predictive models sensitive to changes in microhabitat structure and composition. / Master of Science