Ecosystems transition quickly in the Anthropocene, whereas biodiversity adapts more slowly. Here we simulated a shifting woodland ecosystem on the Colorado Plateau of western North America by using as its proxy over space and time the fundamental niche of the Arizona black rattlesnake (Crotalus cerberus). We found an expansive (= end-of-Pleistocene) range that contracted sharply (= present), but is blocked topographically by Grand Canyon/Colorado River as it shifts predictably northwestward under moderate climate change (= 2080). Vulnerability to contemporary wildfire was quantified from available records, with forested area reduced more than 27% over 13 years. Both 'ecosystem metrics' underscore how climate and wildfire are rapidly converting the Plateau ecosystem into novel habitat. To gauge potential effects on C. cerberus, we derived a series of relevant 'conservation metrics' (i.e. genetic variability, dispersal capacity, effective population size) by sequencing 118 individuals across 846 bp of mitochondrial (mt)DNA-ATPase8/6. We identified five significantly different clades (net sequence divergence = 2.2%) isolated by drainage/topography, with low dispersal (F-ST = 0.82) and small sizes (2N(ef) = 5.2). Our compiled metrics (i.e. small-populations, topographic-isolation, low-dispersal versus conserved-niche, vulnerable-ecosystem, dispersal barriers) underscore the susceptibility of this woodland specialist to a climate and wildfire tandem. We offer adaptive management scenarios that may counterbalance these metrics and avoid the extirpation of this and other highly specialized, relictual woodland clades.

Doctor of Philosophy / Department of Biology / Keith B. Gido / A fundamental goal of ecology is to understand environmental associations of species. These associations can provide a basis for predicting spatial distributions in contemporary habitats as well as how those distributions might change in response to anthropogenic environmental change. Developing species distribution models is limited by an incomplete understanding of functional traits, spatial scaling, and the mechanisms and generalities of correlations among abundance and environmental gradients. I address these four issues using observational and experimental approaches. First, I tested opposing mechanisms of community assembly by measuring the dispersion (i.e., diversity) of three types of functional strategies at three spatial scales and along environmental gradients. I found that communities are assembled via abiotic environmental filtering, but the strength of this filtering depends on the spatial scale of investigation, longitudinal network position, and type of functional strategy. Second, I quantified community-environment relationships across thirteen sub-basins, nested within the three major basins within Kansas to evaluate the consistency (i.e., generality) in predictive capability of environmental variables among sub-basins and across spatial extents. I found that longitudinal network position is consistently the strongest predictor of community composition among sub-basins, but in-stream and catchment predictors become stronger correlates of community composition with increasing spatial extent. Third, I used environmental niche models to quantify distributions of four pairs of congeneric cyprinids and found that species within each pair exhibited contrasting stream-size preferences. I then used field experiments to test for differences in individual-level performance between one pair of species (Pimephales notatus and P. vigilax) along a gradient of stream size. I found that adult spawn success and juvenile growth and condition increased with stream size for both species, indicating that these congeners respond similarly to abiotic gradients associated with the river continuum. I concluded that complementary distributions are a consequence of biotic interactions, differential environmental filtering evident in an unmeasured performance metric, or differential environmental filtering by an environmental factor operating at longer timescales. These studies demonstrate the context dependencies of characterizing habitat associations of stream fishes, but also reveal the general importance of stream size and associated environmental gradients in structuring stream fish communities.

Assessment of the impact of landscape structure on the dispersion of lynx in Posumavi Abstract: The big predators are a traditional group of bioindicative species, whose presence shows the state of environment and landscape within a region. As these animals recolonize the cultural landscape of central Europe it's needed to understand their space requirements and habitat preferences. The lynx (Lynx lynx) has been one of the Sumava forest species since the 70's. After almost 40 years the population of lynx has spread from Sumava and Bavarian Forest to forested parts of Cesky Les, Smrcina and Novohradske hory and to both sides of state borders. However today's state of lynx population, stable number of animals or genetic variability is endangered by many factors. Long term viability of the lynx population is - besides poaching or illegal hunting − affected by the loss of suitable habitats and ongoing fragmentation of the landscape. One of the possible solutions of negative impacts is delineation and protection of landscape permeability of habitat corridors. They might have provide supply of new individuals to small, extinction prone population and at the same time they could reinforce overall fitness population. This thesis has three main objectives:1) evaluation of landscape fragmentation and connectivity in...

京都大学 / 新制・課程博士 / 博士(理学) / 甲第24826号 / 理博第4975号 / 新制||理||1710(附属図書館) / 京都大学大学院理学研究科生物科学専攻 / (主査)准教授 中野, 智之, 教授 朝倉, 彰, 教授 森, 哲 / 学位規則第4条第1項該当 / Doctor of Science / Kyoto University / DGAM

Modelos de distribuição de espécies, também conhecidos como modelos bioclimáticos ou modelos de nicho ecológico, têm sido aplicados em inúmeras questões ecológicas, incluindo espécies invasoras. Essa tese identificou as principais tendências e lacunas de estudos sobre o uso de modelos de distribuição de espécies para predizer a distribuição potencial de espécies invasoras (primeiro artigo). Adicionalmente, aplicou-se modelos de distribuição de espécies para predizer a distribuição potencial de Eragrostis plana Nees na América do Sul e verificar se ocorreu mudança de seu nicho bioclimático durante o processo de invasão (segundo artigo). Finalmente, avaliou-se a resposta em relação às áreas de ocorrência de cinco gramíneas Africanas invasoras nas Américas frente às mudanças climáticas (terceiro artigo). O primeiro artigo realiza uma análise cienciométrica sobre o uso de modelos de distribuição de espécies para predizer a distribuição potencial de espécies invasoras. O segundo artigo utiliza o Algoritmo GARP como técnica de modelagem e foram criados dois modelos para predizer a distribuição potencial de E. plana: um utilizando dados da região nativa da espécie (África do Sul) e outro com dados da região nativa e invadida (regiões da Argentina, Brasil e Uruguai). Posteriormente, cada modelo foi projetado na América do Sul para identificar regiões favoráveis ao estabelecimento de E. plana e verificar se os registros de ocorrência da espécie encontram-se dentro das regiões preditas com alta probabilidade pelos modelos. Além disso, a hipótese de que espécies podem alterar o seu nicho climático durante o processo de invasão foi avaliada para E. plana através de análises estatísticas multivariadas. O terceiro artigo aplica distintas técnicas de modelagem disponíveis no ambiente computacional BIOMOD, seguidas de conjunto de previsões para predizer a distribuição potencial das cinco gramíneas invasoras Africanas nas Américas frente às mudanças climáticas globais. / Species distribution models, also known as bioclimatic models or ecological niche models, have been applied in numerous ecological issues, including invasive species. This thesis indentified the main trends and gaps in studies on the use of species distribution models to predict the potential distribution of invasive species (first paper). Additionally, it includes species distribution modelling to predict the potential distribution of Eragrostis plana Nees in South America and assess the potential shift of its bioclimatic niche in the process of invasion (second paper). Finally, it includes as assessment of the response in terms of areas of occurrence of five invasive African grasses in the Americas under climate changes (third paper). The first paper provides a scientometric analysis on the use of species-distribution models to predict the potential distribution of invasive species. The second paper uses the algorithm GARP as modelling method and created two models to predict the potential distribution of E. plana: the first one used data from the native region (South Africa) and the second one data from both the native and invaded (Argentina, Brazil, and Uruguay). Subsequently, each model was projected in South America to identify regions favorable to the establishment of E. plana and assess whether the occurrence records are found within regions predicted by the models with high probability. Moreover, the hypothesis that species can shift their bioclimatic niche during the invasion process was evaluated for E. plana using multivariate statistical analysis. The third paper applies distinct modelling methods available in the BIOMOD package, followed by ensembles forecasting to predict the potential distribution of five invasive African grasses in Americas under climate changes.

Modelos de distribuição de espécies, também conhecidos como modelos bioclimáticos ou modelos de nicho ecológico, têm sido aplicados em inúmeras questões ecológicas, incluindo espécies invasoras. Essa tese identificou as principais tendências e lacunas de estudos sobre o uso de modelos de distribuição de espécies para predizer a distribuição potencial de espécies invasoras (primeiro artigo). Adicionalmente, aplicou-se modelos de distribuição de espécies para predizer a distribuição potencial de Eragrostis plana Nees na América do Sul e verificar se ocorreu mudança de seu nicho bioclimático durante o processo de invasão (segundo artigo). Finalmente, avaliou-se a resposta em relação às áreas de ocorrência de cinco gramíneas Africanas invasoras nas Américas frente às mudanças climáticas (terceiro artigo). O primeiro artigo realiza uma análise cienciométrica sobre o uso de modelos de distribuição de espécies para predizer a distribuição potencial de espécies invasoras. O segundo artigo utiliza o Algoritmo GARP como técnica de modelagem e foram criados dois modelos para predizer a distribuição potencial de E. plana: um utilizando dados da região nativa da espécie (África do Sul) e outro com dados da região nativa e invadida (regiões da Argentina, Brasil e Uruguai). Posteriormente, cada modelo foi projetado na América do Sul para identificar regiões favoráveis ao estabelecimento de E. plana e verificar se os registros de ocorrência da espécie encontram-se dentro das regiões preditas com alta probabilidade pelos modelos. Além disso, a hipótese de que espécies podem alterar o seu nicho climático durante o processo de invasão foi avaliada para E. plana através de análises estatísticas multivariadas. O terceiro artigo aplica distintas técnicas de modelagem disponíveis no ambiente computacional BIOMOD, seguidas de conjunto de previsões para predizer a distribuição potencial das cinco gramíneas invasoras Africanas nas Américas frente às mudanças climáticas globais. / Species distribution models, also known as bioclimatic models or ecological niche models, have been applied in numerous ecological issues, including invasive species. This thesis indentified the main trends and gaps in studies on the use of species distribution models to predict the potential distribution of invasive species (first paper). Additionally, it includes species distribution modelling to predict the potential distribution of Eragrostis plana Nees in South America and assess the potential shift of its bioclimatic niche in the process of invasion (second paper). Finally, it includes as assessment of the response in terms of areas of occurrence of five invasive African grasses in the Americas under climate changes (third paper). The first paper provides a scientometric analysis on the use of species-distribution models to predict the potential distribution of invasive species. The second paper uses the algorithm GARP as modelling method and created two models to predict the potential distribution of E. plana: the first one used data from the native region (South Africa) and the second one data from both the native and invaded (Argentina, Brazil, and Uruguay). Subsequently, each model was projected in South America to identify regions favorable to the establishment of E. plana and assess whether the occurrence records are found within regions predicted by the models with high probability. Moreover, the hypothesis that species can shift their bioclimatic niche during the invasion process was evaluated for E. plana using multivariate statistical analysis. The third paper applies distinct modelling methods available in the BIOMOD package, followed by ensembles forecasting to predict the potential distribution of five invasive African grasses in Americas under climate changes.

Modelos de distribuição de espécies, também conhecidos como modelos bioclimáticos ou modelos de nicho ecológico, têm sido aplicados em inúmeras questões ecológicas, incluindo espécies invasoras. Essa tese identificou as principais tendências e lacunas de estudos sobre o uso de modelos de distribuição de espécies para predizer a distribuição potencial de espécies invasoras (primeiro artigo). Adicionalmente, aplicou-se modelos de distribuição de espécies para predizer a distribuição potencial de Eragrostis plana Nees na América do Sul e verificar se ocorreu mudança de seu nicho bioclimático durante o processo de invasão (segundo artigo). Finalmente, avaliou-se a resposta em relação às áreas de ocorrência de cinco gramíneas Africanas invasoras nas Américas frente às mudanças climáticas (terceiro artigo). O primeiro artigo realiza uma análise cienciométrica sobre o uso de modelos de distribuição de espécies para predizer a distribuição potencial de espécies invasoras. O segundo artigo utiliza o Algoritmo GARP como técnica de modelagem e foram criados dois modelos para predizer a distribuição potencial de E. plana: um utilizando dados da região nativa da espécie (África do Sul) e outro com dados da região nativa e invadida (regiões da Argentina, Brasil e Uruguai). Posteriormente, cada modelo foi projetado na América do Sul para identificar regiões favoráveis ao estabelecimento de E. plana e verificar se os registros de ocorrência da espécie encontram-se dentro das regiões preditas com alta probabilidade pelos modelos. Além disso, a hipótese de que espécies podem alterar o seu nicho climático durante o processo de invasão foi avaliada para E. plana através de análises estatísticas multivariadas. O terceiro artigo aplica distintas técnicas de modelagem disponíveis no ambiente computacional BIOMOD, seguidas de conjunto de previsões para predizer a distribuição potencial das cinco gramíneas invasoras Africanas nas Américas frente às mudanças climáticas globais. / Species distribution models, also known as bioclimatic models or ecological niche models, have been applied in numerous ecological issues, including invasive species. This thesis indentified the main trends and gaps in studies on the use of species distribution models to predict the potential distribution of invasive species (first paper). Additionally, it includes species distribution modelling to predict the potential distribution of Eragrostis plana Nees in South America and assess the potential shift of its bioclimatic niche in the process of invasion (second paper). Finally, it includes as assessment of the response in terms of areas of occurrence of five invasive African grasses in the Americas under climate changes (third paper). The first paper provides a scientometric analysis on the use of species-distribution models to predict the potential distribution of invasive species. The second paper uses the algorithm GARP as modelling method and created two models to predict the potential distribution of E. plana: the first one used data from the native region (South Africa) and the second one data from both the native and invaded (Argentina, Brazil, and Uruguay). Subsequently, each model was projected in South America to identify regions favorable to the establishment of E. plana and assess whether the occurrence records are found within regions predicted by the models with high probability. Moreover, the hypothesis that species can shift their bioclimatic niche during the invasion process was evaluated for E. plana using multivariate statistical analysis. The third paper applies distinct modelling methods available in the BIOMOD package, followed by ensembles forecasting to predict the potential distribution of five invasive African grasses in Americas under climate changes.

La connaissance des facteurs qui influent sur les populations de tiques et en particulier sur leurs distributions est un préalable indispensable à l’étude des maladies qu’elles transmettent. Parmi eux, l’importance des facteurs biotiques et en particulier celle de la compétition interspécifique est peu connue et souvent négligée. L’objectif de ce travail était d’évaluer le rôle de la compétition interspécifique sur la distribution de deux espèces de tiques d’importance vétérinaire, Amblyomma variegatum et A. hebraeum. Alors que seule A. variegatum favorise la transmission de la dermatophilose, une maladie de peau débilitante pour les ruminants, les deux espèces sont vectrices de la cowdriose, maladie due à une bactérie Ehrlichia ruminantium qui représente une contrainte économique majeure pour l’élevage. L’impact de l’espèce vectrice dans l’épidémiologie de la cowdriose (différences de souches circulantes, sévérité des cas,…) est inconnu bien qu’ayant potentiellement des conséquences en termes de surveillance (risque épidémiologique à l’introduction d’animaux) et de contrôle (développement de vaccin régional adapté aux souches circulantes) de la maladie. Ces deux espèces ont une distribution contiguë en Afrique australe avec peu de chevauchement (distribution parapatrique) suggérant une préférence environnementale différentielle ou une compétition exclusive entre elles. Une revue des données de la littérature a permis de mettre en évidence un chevauchement de leur niche trophique, climatique, et temporelle, et une interférence comportementale via la production de leurs phéromones. Les deux espèces pourraient donc rentrer en compétition directement par interférence communicative ou reproductive (hybridation stérile), ou indirectement via le partage de ressources, prédateurs ou pathogènes communs. Le rôle des facteurs biotiques et abiotiques sur le maintien de la parapatrie de ces deux espèces a été analysée (i) d’une part par la comparaison de leur niche environnementale réalisée en Afrique australe, et aux deux zones de contact au Zimbabwe et Mozambique, par des méthodes d’ordination et de modèles de niche (Maxent) ; (ii) et d’autre part par l’étude de leurs distributions et de leurs interactions (distribution sur les hôtes, croisements hétérospécifiques) dans la zone de contact au Mozambique. Globalement les résultats montrent qu’en Afrique Sud-Est et au Zimbabwe les deux espèces occupent des niches environnementales distinctes, celle d’A. hebraeum incluant des zones plus chaudes et plus sèches que celle d’A. variegatum. Au contraire au Mozambique les deux niches se chevauchent considérablement. L’enquête de terrain dans cette zone montre que les deux espèces y sont moins souvent trouvées en sympatrie que les données prédites, suggérant une distribution en partie déterminée par des interactions biotiques. Dans les rares sites avec présence des deux espèces, A. variegatum et A. hebraeum partagent les mêmes sites de fixation sur les animaux et forment un pourcentage relativement élevé de couples hétérospécifiques. Ce pourcentage, bien que similaire entre les femelles A. variegatum et A. hebraeum, semble impliquer des processus de discrimination spécifique propres aux deux espèces intervenant au niveau de l’agrégation, de la fixation et du contact des individus. Nos résultats suggèrent l’existence d’une compétition exclusive entre les deux espèces, due à une compétition sexuelle probablement associée à une compétition indirecte. La frontière parapatrique semble occuper une position stable le long d’un gradient environnemental au Zimbabwe mais pas au Mozambique. Les conditions entrainant la coexistence ou l’exclusion des deux espèces avec formation d’une frontière parapatrique sont discutés à l’aide de modèles théoriques de compétition. / Studying the factors that influence tick populations and their distributions is an essential pre-requisite to understanding the dynamics of the diseases they transmit. The relative importance of biotic factors such as interspecific competition is not well known and often neglected. The objective of this study was to assess the influence of interspecific competition on the distribution of two tick species of veterinary importance, Amblyomma variegatum and A. hebraeum. Whereas only A. variegatum is known to favor dermatophilosis, a debilitating skin disease of ruminants, both species are good vectors of Ehrlichia ruminantium, the bacteria causing heartwater, a fatal disease of ruminants that presents a major constraint for livestock development in Africa. The impact of vector species in heartwater epidemiology (differences of circulating strains, severity of clinical cases…) is poorly known but may have important implications for surveillance (epidemiological risk of imported animals) and control (adapting regional vaccine programs to circulating strains) of the disease. These two ticks have abutting and marginally overlapping (i.e. parapatric) distributions in southern Africa, suggesting either differential environmental preferences or exclusive competition between the two species. A review of published data highlighted an important overlap of their trophic, climatic and seasonal niche, and existence of chemical behavior interference through pheromone production. Therefore, the two species might compete either directly by communicative or reproductive interference (sterile hybridization), or indirectly by sharing the same resources, predators or pathogens. The role of biotic and abiotic factors in determining parapatry of these species was assessed by (i) comparing their realized environmental niche in southern Africa, and at contact zones in Zimbabwe and Mozambique, using ordination techniques and environmental niche modeling (Maxent); (ii) studying their distributions and their interactions (distribution on co-infested host, heterospecific mating) at the contact zone in Mozambique. Globally, the results indicated the two species occupied distinctly different environmental niches in southern Africa and at the contact zone in Zimbabwe, with the niche of A. hebraeum including both hotter and drier areas than that of A. variegatum. However, in Mozambique their niches overlapped considerably. Field studies within this zone showed that sympatry was observed less frequently than predicted by niche models, suggesting an importance of biotic interactions. At the rare sites where both species were present, A. variegatum and A. hebraeum were observed to share the same preference sites on hosts and formed a high percentage of heterospecific pairs. Though this cross-mating rate was not significantly different between A. variegatum and A. hebraeum females, our observations suggest different mechanism of species discrimination involved for the two species acting at the aggregation, fixation and partner contact level. Our results strongly suggest exclusive competition between these species may arise from sexual competition probably interacting with other indirect forms of competition. The parapatric boundary apparently occupies a stable location along an environmental gradient at the contact zone in Zimbabwe but not in Mozambique. Conditions inducing coexistence or exclusion of both species with the formation of parapatric distributions are discussed in relation to theoretical models of competition. When sexual competition is introduced in these models, the relative frequency of two species is determined by their endogenous fitness (a function of environmental conditions), density dependent effects of competitive interactions, historical distributions and dispersal rates: survival of the first prevails when the immigration rate of a fitter invading species remains lower than an invasion threshold.

Ecological niche modelling is a method designed to describe and predict the geographic distribution of an organism. This procedure aims to quantify the species-environment relationship by describing the association between the organism’s occurrence records and the environmental characteristics at these points. More simply, these models attempt to capture the ecological niche that a particular organism occupies. A popular application of ecological niche models is to predict the potential distribution of invasive alien species in their introduced range. From a biodiversity conservation perspective, a pro-active approach to the management of invasions would be to predict the potential distribution of the species so that areas susceptible to invasion can be identified. The performance of ecological niche models and the accuracy of the potential range predictions depend on the quality of the data that is used to calibrate and evaluate the models. Three different types of input data can be used to calibrate models when producing potential distribution predictions in the introduced range of an invasive alien species. Models can be calibrated with native range occurrence records, introduced range occurrence records or a combination of records from both ranges. However, native range occurrence records might suffer from geographical bias as a result of biased sampling or incomplete sampling. When occurrence records are geographically biased, the underlying environmental gradients in which a species can persist are unlikely to be fully sampled, which could result in an underestimation of the potential distribution of the species in the introduced range. I investigated the impact of geographical bias in native range occurrence records on the performance of ecological niche models for 19 invasive plant species by simulating two geographical bias scenarios (six different treatments) in the native range occurrence records of the species. The geographical bias simulated in this study was sufficient to result in significant environmental bias across treatments, but despite this I did not find a significant effect on model performance. However, this finding was perhaps influenced by the quality of the testing dataset and therefore one should be wary of the possible effects of geographical bias when calibrating models with native range occurrence records or combinations there of. Secondly, models can be calibrated with records obtained from the introduced range of a species. However, when calibrating models with records from the introduced range, uncertainties in terms of the equilibrium status and introduction history could influence data quality and thus model performance. A species that has recently been introduced to a new region is unlikely to be in equilibrium with the environment as insufficient time will have elapsed to allow it to disperse to suitable areas, therefore the occurrence records available would be unlikely to capture its full environmental niche and therefore underestimate the species’ potential distribution. I compared model performance for seven invasive alien plant species with different simulated introduction histories when calibrated with native range records, introduced range records or a combination of records from both ranges. A single introduction, multiple introduction and well established scenario was simulated from the introduced range records available for a species. Model performance was not significantly different when compared between models that were calibrated with datasets representing these three types of input data under a simulated single introduction or multiple introduction scenario, indicating that these datasets probably described enough of the species environmental niche to be able to make accurate predictions. However, model performance was significantly different for models calibrated with introduced range records and a combination of records from both ranges under the well established scenario. Further research is recommended to fully understand the effects of introduction history on the niche of the species. Copyright / Dissertation (MSc)--University of Pretoria, 2009. / Zoology and Entomology / unrestricted

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