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.

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

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.