Abstract In this thesis, I studied spatial and temporal variability of macroinvertebrate assemblages of boreal streams. The main objectives were (i) to characterize macroinvertebrate assemblage types across large geographical extents, and to assess the utility of assemblage types and landscape and stream type classifications as the basis of stream bioassessment and conservation programs (ii, iii). I also examined the relative roles of large-scale spatial trends and local environmental conditions in structuring macroinvertebrate assemblages (iv). Finally, I assessed (v) if assemblage classifications produce temporally stable and predictable assemblage types. Stream macroinvertebrate assemblage structure exhibited continuous variation instead of distinct assemblage types. Although ecoregions clearly accounted for a considerable amount of variation of macroinvertebrate assemblage characteristics, a combination of regional stratification and prediction from environmental factors would probably yield the most comprehensive framework for the characterizations of macroinvertebrate assemblages of boreal headwater streams. Differences in macroinvertebrate assemblage structure, as well as a group of effective indicator species for different stream types, suggest that landscape classifications could be used as a preliminary scheme for the conservation planning of running waters The strength of the relationship between assemblage structure and local environmental variables increased with decreasing extent, whereas assemblage variation related to spatially variables showed the opposite pattern. At the largest scale, spatial variation was related to latitudinal gradients, while spatial autocorrelation among neighbouring streams was the likely mechanism creating spatial structure within drainage systems. These results suggest that stream bioassessment should give due attention to spatial structuring of stream assemblage composition, considering that important assemblage gradients may not only be related to local environmental factors, but also to biogeographical constraints and neighbourhood dispersal processes. The classification strengths of macroinvertebrate assemblages based on data of three years were overall rather weak, and more importantly, the compositions of the site groups varied considerably from year to year. Such wide and continuous variation was also mirrored by low and inconsistent predictability of classifications from environmental variables. The observed level of temporal variation in assemblage structure may not be a serious problem for predictive approaches frequently used in bioassessment of freshwater ecosystems. For conservation purposes, however, alternative approaches (e.g. physical surrogates of biodiversity) need to be considered.

Human activities have transformed natural landscapes into human-dominated areas at unprecedented rates in the last centuries. Land cover transformation is associated with loss of natural habitat, thus a threat to biodiversity. Because habitat loss will likely continue in the future due to population growth and increase demand for natural resources, an important question in ecological studies is whether land cover features (i.e. amount, variety, shape, configuration) can be used as predictors to estimate species loss from habitat modification. This thesis investigates the predictive ability of landscape features in predicting species distributions at the landscape level and across large regions. It tests several predictions from classic hypotheses such as the species-area relationship and habitat fragmentation, utilizing a macroecological approach. Response variables (e.g. species richness, species’ probability of occurrence) and independent variables (e.g. proportion of natural areas, metrics of fragmentation, temperature, etc.) are analysed in cell sizes of 25-900km2 covering large regions (e.g. southern Ontario, New York State). Bird species were chosen as the main biological model. Most literature assumes that species richness should vary positively as a function of remaining natural area, following the well-known species–area relationship (i.e. classic SAR). Prior studies have shown that avian species richness has a peaked, rather than a monotonic increasing, relationship with the proportion of natural land cover in landscapes of southern Ontario. The first chapter of the thesis showed improvements in the predictive power of classic SARs by proposing the “Lost-habitat SAR”, which demonstrates that richness of open-habitat species can be predicted when we partition human-dominated land cover into an ‘‘available human-dominated’’ component and ‘‘lost’’ habitat (i.e. parts of the landscape that can no longer be utilized by any species). The second chapter addresses a current contention in the literature about the effect of habitat fragmentation beyond habitat amount at the landscape level. Specifically, I tested the effect of fragmentation (e.g. number of patches) on both avian richness and the probability of occurrence (pocc) of individual species, after controlling for habitat amount in 991 landscapes, each 100-km2, in southern Ontario. The analysis showed that overall species richness responds primarily to habitat amount, and that the effect of habitat fragmentation, holding the total amount of habitat constant, is negligible. The probability of occurrence of a few bird species did relate negatively to the size, number and isolation of the patches within the landscape. We argue that the evidence is inconsistent with the hypothesis that reducing habitat fragmentation would be an effective conservation strategy for birds at the landscape level. Chapter 3 tested the speculation in the climate change literature that habitat loss may impede the colonization or movement of species whose ranges are shifting northwards in response to climate. Using the same 100-km2 landscapes of southern Ontario, I examined individual bird species’ probability of occupancy as a function of the amount of remaining natural land cover for three groups of species: i) those whose northern range limit falls within the study area, ii) those whose southern range limit is in the study area, and iii) those whose ranges cover the entire study area. The results showed that the probability of occupancy of southern-edge species is a positive function of the amount of natural land cover (forest) in the landscape, while the probability of occupancy of northern-edge species is a negative function of natural land cover. Hence, I conclude that at southern range limits species faces the dual stresses of climatic warming and habitat conversion. Whereas, at northern (potentially expanding) range edges, partially disturbed landscapes are more readily occupied than undisturbed landscapes. In the final chapter, I challenge widely accepted hypothesis that habitat loss causes biodiversity loss by testing whether conserving natural land cover would conserve species diversity. More specifically, I tested whether broad-extent relationships between avian species richness and natural land cover are independent of: 1) whether species distribution data come from systematic censuses (atlases) versus range maps, and 2) the grain size of the analysis in grid cells covering southern Ontario, CA, and New York State, US. My findings showed that over regional extents, range-map-based richness relates strongly to temperature, irrespective of spatial grain, and that censused species richness relates to temperature less strongly. Moreover, range-map richness is a negative function of the proportion of natural land cover, while realized richness is a peaked function. Therefore, I conclude that conserving natural land cover would not conserve species diversity in southern Ontario or in New York State, since greater natural cover does not imply higher richness. We argue that habitat loss has become a panchreston. It may be misguiding conservation biology strategies by focusing on a threat that is too general to be usefully predictive.

Low light intensity habitats harbor unique sciophilous benthic communities and are a source of novel and unique sponge fauna. However, the community structure of these habitats is poorly studied to date. Thus, this study attempts to understand the composition and structure of sciophilous sponge populations in southeast Florida. Fifty limestone plates were placed on a shallow reef in Fort Lauderdale for two years (2010-2012). To identify the sponge community and their patterns over time, all plates were photographed at the end of each year. Then, samples were taken from each of the live sponge specimens observed on the plates and processed in the laboratory for taxonomical identification. A total of 45 different sponge species were found, the majority corresponding to the Poecilosclerida Order. Eighteen were identified to species level, twenty-two to genus, and five were undetermined. Eight sponges constitute new records to Florida, and four are potentially new species. The most dominant species include Oscarella sp.1, Dysidea etheria, Mycale sp.1, Halisarca caerula and Tedania ignis. Species richness significant varied among years, and species cover among sectors (inner and outer reef). However, sponge assemblages were similar between years with slightly variation between sectors. This study found a diverse and complex composition and structure of sponges that is quite distinct from the sponge assemblages on the open reefs. A guide of the biodiversity of cryptic sponge species was created to facilitate further studies in low light intensity habitats.

Abstract Recognition of the importance of a regional perspective for understanding the structure and dynamics of local assemblages has stimulated the emergence of the field of macroecology. Most attention has been directed to terrestrial ecosystems, while large-scale patterns in biodiversity of aquatic organisms have received less attention. In this thesis I examined patterns of aquatic diversity across several geographic areas and scales, in an effort to understand some of the environmental and spatial factors determining species diversity in aquatic environments. The main objectives of this thesis were: (i) to examine the latitudinal diversity patterns of marine crustaceans and molluscs and their relationship to large scale environmental gradients, (ii) to study macroinvertebrate species richness in headwater streams at two spatial extents, within and across drainage systems, and assess the relative importance of local, landscape and regional variables, and (iii) to study diversity patterns of macroorganisms vs microorganism, comparing distance decay patterns of stream diatoms, macroinvertebrates and bryophytes. Latitudinal diversity patterns of crustaceans and molluscs were clearly related to larval developmental mode. An increase in species richness towards high latitudes was found for species with direct development, whereas richness of species with planktotrophic development decreased poleward. Sea surface temperature was the most important environmental gradient related to species richness of both phyla and each developmental mode, but with different effects on each mode. Stream macroinvertebrate species richness at the bioregion extent was negatively related to water humic content. Another factor related to species richness at the bioregion extent was elevation range, a variable linked to stream topographic heterogeneity. Local environmental variables explained most of the variation in species richness at the drainage system extent, however high among-region variability was evident. Patterns between macro- and microorganism may not be fundamentally different, but the level of environmental control varied, being strongest for diatoms, while some groups of benthic macroinvertebrates exhibited relatively strong dispersal limitation. The relative importance of niche vs. dispersal processes is not simply a function of organism size but other traits (e.g. life-history type, dispersal capacity) may obscure this relationship.

Developing consistent and repeatable broad-scale methods for biodiversity modelling is an important goal to address as habitat loss, fragmentation and environmental degradation threaten our ability to maintain ecosystem and species diversity levels. Geospatial reviews of biodiversity monitoring have identified ecological indicators for the indirect mapping of species richness and ecosystem components modelling the processes controlling species distribution gradients. The goal of our research is to advance broad-scale biomonitoring by demonstrating how landscape-scale environmental indices can be used to model regional ecosystem and species diversity of British Columbia (BC), Canada. We meet our ecosystem-modelling goal by selecting and developing suitable ecological indicators from Earth observation data and terrain indices to represent the structure, composition and function of the environment, displaying both static and dynamic landscape processes of BC’s ecosystems. We regionalize the selected indirect indicators of biodiversity using a two-step clustering algorithm. The results display 16 ecologically distinct terrestrial ecosystems, 10 of which characterize the northern Boreal, coastal and Southern Interior mountain regions, and six represent the coastal lowlands, interior, Georgia Depression, Boreal and Taiga Plains of British Columbia. Comparing our classification to BC Ministry of Forests biogeoclimatic zone mapping, we find spatial similarity in the coastal, Taiga and Boreal Plains. Overall, our classification distinguishes a greater diversity of ecosystems in the mountainous regions of the province and greater homogeneity in the Central Interior where our landscape characteristics represent current productivity conditions. Our approach to ecosystem modelling supports legacy mapping by providing ecological information in under-sampled regions of BC and offers a method for consistent repeat modelling of ecosystem diversity to identify landscape change. To meet our species-modelling goal we employ a flexible non-parametric regression tree model (Random Forests) to establish the power of landscape-scale indicators (productivity, ambient energy, and heterogeneity) to predict the spatial distribution of breeding bird richness and establish the dominant landscape processes controlling vertebrate richness throughout BC. Our models explain approximately 40% of the variation in survey effort stratified breeding bird species richness levels and distinguish ambient energy as the top ranked environmental predictors of breeding richness. Using our modelled relationships, we forecast breeding richness levels for the regions of BC not currently surveyed to support conservation management of birds and vertebrate species. The results identify the lowland, warm and dry regions of the Boreal, Taiga, South and Central Interior and the Georgia Depression to be species rich. These results have implications for conservation managers, as high breeding richness is also concentrated in the areas favourable to human settlement. Additionally, by connecting breeding bird data derived from remotely sensed data and continuously collected climate data, we provide an approach for monitoring ecological indicators as surrogates of vertebrate population levels over broad spatial scales. / Graduate

Magister Scientiae (Biodiversity and Conservation Biology) / Since the late 1970 there has been increased concern of amphibian decline and extinction. Several causes for the worldwide declines have been suggested and include ultraviolet radiation, predation, pollution, climate change, diseases and habitat modification. To counter this, more research on the subject has been encouraged of which long term monitoring has been suggested as a research method. The study was conducted in Kakamega Forest in Kenya, which is the country's remnant of the once vast Guineo-Congolian forest. A rectangular transect whose sides measured 600 m in total was established and transect walks were carried out every two weeks for two consecutive days between 2002 and 2006. 24 species were targeted in the study and were sampled through VES and AES and data recorded in a GPS and later downloaded. In this study I examined the influence of rainfall, temperature, habitat and moon phases on the activity of frogs in Kakamega Forest. I also determined under which weather conditions sampling was more efficient. When monitoring was carried out by two observers I tested whether their data were similar. Data were analysed using non-parametric methods (Kruskal-wallis and Tukey test), species abundances analysed using EstimateS..Out of the 24 targeted species only 14 were recorded, with a total of 535 specimens being counted mostly at night. Most frogs in Kakamega Forest were more active in temperatures between 20 and 25oC. There was not much variation and there was no frog activity when the temperature was extremely high. There was rainfall throughout the year and there was no significant differences in the number of frogs counted in rainfall above 200 mm or below 200 mm. There was no significant difference in the number of specimens found in the different vegetation segments in the forest. More amphibians were caught under cloudy, rainy and clear conditions at night than under any weather condition during the day. During the day, more amphibians were caught during cloudy conditions than when it rained or when there was no cloud cover. There was no difference in catch among night conditions and there was no difference between clear and rainy days In Kakamega Forest, night is the best time to sample amphibians. In terms of weather it is best to sample when it is cloudy both during the day and at night. There were no differences in sampling abilities between two observers tested under similar weather conditions.

I studied the spatial distribution of South African avian species richness from the viewpoint that humans are a substantial modifying force on earth, and have also modified the historical spatial distribution of species richness. The main aim of the thesis is to investigate the way in which humans have modified avian species richness patterns in South Africa at the quarter-degree square (QDS) resolution, which is a phenomenon that has been either overlooked, or not completely clarified, in many previous studies of the same region and data at the same resolution. In particular, I investigated hypotheses that were proposed to explain the maintenance of a positive relationship between native species richness and human population density in the face of negative human impacts. Further, I investigated which of the possible anthropogenic and natural environmental factors determine spatial distribution in exotic bird species. Highlighted from these studies are that substantial positive and negative human influences on bird species richness distribution patterns are observable at the QDS resolution, that there are differences between common native birds and rare native birds with regard to their relationships with anthropogenic environmental conditions and exotic bird species, and that the particular combination of environmental covariates that is important for the spatial distributions of exotic species is taxon- and scale-dependent. Even though these results have contributed much towards our understanding on how human modifications have affected species richness patterns, this thesis leaves some unanswered questions. Finer resolution studies and temporal studies are needed to examine many of these questions. Further, an interdisciplinary approach incorporating politics and economics into ecological studies is needed to enhance our understanding of the factors that modify the distribution of humans and their associated threats and benefits to species richness. / Dissertation (MSc)--University of Pretoria, 2010. / Zoology and Entomology / unrestricted

The species–area relationship (SAR) has been applied to predict species richness declines as area is converted to human-dominated land covers.In many areas of the world, however, many species persist in human-dominated areas, including threatened species. Because SARs are decelerating nonlinear, small extents of natural habitat can be converted to human use with little expected loss of associated species, but with the addition of more species that are associated with human land uses. Decelerating SARs suggest that, as area is converted to human-dominated forms, more species will be added to the rare habitat than are lost from the common one. This should lead to a peaked relationship between richness and natural area. I found that the effect of natural area on avian richness across Ontario was consistent with the sum of SARs for natural habitat species and human-dominated habitat species, suggesting that almost half the natural area can be converted to human-dominated forms before richness declines. However, I found that this spatial relationship did not remain consistent through time: bird richness increased when natural cover was removed (up to 4%), irrespective of its original extent. The inclusion of metapopulation processes in predictive models of species presence improves predictions of diversity change through time dramatically. Variability in site occupancy was common among bird species evaluated in this study, likely resulting from local extinction-colonization dynamics. Likelihood of species presence declined when few neighbouring sites were previously occupied by the species. Site occupancy was also less likely when little suitable habitat was present. Consistent with expectations that larger habitats are easier targets for colonists, habitat area was more important for more isolated sites. Accounting for the effect of metapopulation dynamics on site occupancy predicted change in richness better than land cover change and increased the strength of the regional richness–natural area relationship to levels observed for continental richness–environment relationships suggesting that these metapopulation processes “scale up” to modify regional species richness patterns making them more difficult to predict. It is the existence of absences in otherwise suitable habitat within species’ ranges that appears to weaken regional richness–environment relationships.

It is generally assumed that when natural habitat is converted to human-dominated cover such area is “lost” to its native species. Extinctions will ensue. The literature generally assumes that species are extirpated as natural area is reduced, following the well-known species-area relationship (SAR). However, SARs have consistently over-estimated species losses resulting from conversion of natural habitat to human-dominated land covers. We hypothesize that the overestimation occurs because these area-based models assume that converted habitat is “lost”, eliminating all species. However, in the real world, conversion of natural land cover to human-dominated cover frequently produces new land covers, different from the original habitat, but not necessarily completely inhospitable to biodiversity. We evaluated the responses of total avian richness, forest bird richness and open habitat bird richness to remaining natural area within 991 quadrats, each 100 km2, across southern Ontario. Total bird species richness does not follow SAR predictions; rather, the number of bird species peaks at roughly 50% natural land cover. The richness of forest birds does follow the usual SAR power-law as a function of forested area. In contrast, richness of birds that prefer open-habitat does not increase monotonically with either natural- or human-dominated land cover. However, we can partition human-dominated land cover into an “available human-dominated” component and “lost” habitat. Richness of open-habitat species relates to the amount of available human-dominated cover. Distinguishing three habitat types (natural, available human-dominated, and lost) permits accurate predictions of species losses in response to natural habitat conversion.

Dung beetles is a group of coprophagous beetles in the family Scarabaeidae. Dung beetles have an important function by improving nutrient circulation and biological control of parasites. Dung is a short-lived resource, making dung beetles tied to habitats continuously grazed. Decline in pasture combined with shorter grazing seasons has resulted in a decrease in available dung. Other reasons can be the use of chemicals to control internal parasites on the grazing animals. This has contributed to the fact that half of the dung beetles in Sweden are endangered. The aim of the present study was to analyze data on dung beetles from the County Administration Board in Östergötland and see if species communities have changed between 2001-2019. In addition, the effect of sand as soil type was studied by comparing the species richness between pastures with and without sand. Analysis whether there was any variation among species richness and species communities depending of grazing animals was also performed. Results showed that the number of species had increased over time and that changes in species communities had occurred. Pastures with sand had more species, however, the amount of sand in the nearest surroundings was of less importance. Dung beetles were found to have a preference when choosing dung-type. Pastures with dung from multiple species of grazing animals were found having the highest species richness. Studies on this would clarify even further what seems to favor dung beetles, this could then be applied to other areas to counteract the loss of species.