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Institut
Geographic ranges of species and their determinants are of great interest in the field of biogeography and are often studied in terms of the species" ecological niches. In this context, the range of a species is defined by the accessibility of an area, abiotic factors and biotic interactions, which affect a species" distributions with different intensities across spatial scales. Parapatry describes a distributional pattern in which the ranges of two species meet along sharp range limits with narrow contact zones. Such parapatric range limits are determined by changing abiotic conditions along sharp environmental gradients or can result from interspecific resource competition. However, it has been shown that often the interplay of abiotic conditions and species interactions determine parapatry. The geographic ranges of the land salamanders, Salamandra salamandra and S. atra, narrowly overlap in the European Alps with only few syntopic localities and to date, the cause of parapatry is unknown. The goal of this thesis was thus to identify the importance of abiotic and biotic factors for their parapatric range limits at different spatial scales. On a broad spatial scale, the role of climate for the parapatric range limits of the species was investigated within three contact zones in Switzerland. Climatic conditions at species" records were analysed and species distribution modelling techniques were used to explore the species" climatic niches and to quantify the interspecific niche overlap. Furthermore, it was tested whether the parapatric range limit coincides with a strong climatic gradient. The results revealed distinct niches for the species as well as the presence of strong climatic gradients which could explain the parapatric range limits of the species. Yet, there was a moderate interspecific niche overlap in all contact zones indicating that the species may co-occur and interact with each other in areas where they both find adequate conditions. Comparison among contact zones revealed geographic variation in the species" niches as well as in the climatic conditions at their records suggesting that the species can occur in a much wider range of conditions than they actually do. These findings imply that climate represents a main factor for the species" parapatric range limits. Yet, interspecific niche overlap and the geographic variation provide indirect evidence that interspecific interaction may also affect their spatial distribution. To test whether competition restricts the species" ranges on the habitat scale and to understand local syntopic co-occurrence of the salamanders within their contact zones, site-occupancy modelling was used. This approach allowed to find the habitat predictors that best explain the species" local distribution. While the slope of the site positively affected the occupancy probability of S. salamandra, no tested predictor explained that of S. atra. Also, there was no effect of the occurrence of one species on the occupancy probability of the other providing no evidence for competition. Should competition occur, it does not lead to spatial segregation of the species on this scale. Because biotic interactions most significantly affect the ranges of species on small spatial scales, the microhabitat conditions at locations of the species within syntopic contact zones were compared and a null model analysis was applied to determine their niche overlap. Resource selection probability function models were used to assess those attributes that affect the species" habitat selections. The results revealed species-specific microhabitat preferences related to leaf litter cover, tree number and that the species were active at different temperatures as well as times of the day. The high degree of diurnal activity of S. atra may be due to its preference of forest floor microhabitats that long remain suitable during daytime. Besides, there was a great niche overlap for shelters indicating that the species may compete for this resource. Differential habitat selection and the use of the available shelters at different times of the day may minimize species interactions and allow their local co-occurrence within contact zones. To identify whether the potential infection with the pathogenic chytrid fungus could serve as an alternative biotic explanation for the range margins of S. atra, several populations throughout its range were screened for infection. Since the occurrence of this pathogen was detected mostly at lower altitudes of the Alps, it may confine the range of S. atra to higher elevations. Because chytrid was not detected in any of the samples, the pathogen unlikely plays a role in determining its range limits. Overall, these findings underline the complexity of mechanisms that determine the range margins of parapatric species and provide an important basis for subsequent studies regarding the determinants of the parapatric distribution of the two salamander species.
Comparing the results of the phylogeographies of the four species included in this thesis, some accordances have been found, even though certain patterns are only represented in one or two species. In all cases, the findings of the studied species strongly support the existence of forests or forest-like ecosystems beyond the classic forest refugia in the Mediterranean areas (Iberian, Apennine and Balkan peninsulas) during glacial times. However, evidence of glacial refugial areas in Southeastern Europe, especially the Balkans, have been found in this study as well. The analysed populations of Aposeris foetida, Melampyrum sylvaticum and Erebia euryale showed high genetic diversity values and mostly higher private fragments in this area, which is a strong indicator for centres of glacial survival during Würm and, regarding the results of M. sylvaticum, even during the Riss ice age. Three of the analysed species (A. foetida, M. sylvaticum and Colias palaeno) supported a second main glacial refuge area located along the Northern Alps. Again, high genetic diversity values and the uniqueness of the populations living in this region today prove the importance of this area as a glacial centre of survival. Those results confirm several recently published studies on forest species and strongly indicate the persistence of forest-like structures or even forests during the ice ages along the foothills of the Northern Alps. Additionally, the persistence of C. palaeno in this area furthermore supports the existence of peatlands north of the Alps, at least during the last glacial. The results of M. sylvaticum and E. euryale further indicate the vicinity of the Tatra Mountains as core areas for glacial survival. However, the genetic patterns found for E. euryale are ambiguous. Due to an intermediate position of two genetic lineages (originating in the Eastern Alps and Southeastern Europe), the Tatras could also reflect a postglacial mixture zone of those lineages. Moreover, the glacial and postglacial importance of this area for woodland species was accentuated, supporting other phylogeographic studies published. Besides the congruities among the results of the study species, some unique patterns and therefore further potential glacial refugia have also been illuminated in this thesis. For instance, the calcicole species, A. foetida, most probably had further survival area at both sides of the Dinaric Alps, supported by high genetic diversity values and a high number of private fragments found in Croatian populations. Furthermore, the surroundings of the German Uplands and the margin of the Southern Alps provided suitable conditions for glacial survival for M. sylvaticum, while the Eastern and Southeastern Alpine region most probably sheltered the Large Ringlet E. euryale during ice ages. Additionally, this butterfly species survived at least the glaciation along the foothills of the Massif Central, whose present populations showed a unique genetic lineage and their genetic diversity values have been measurably higher than in other populations for this species. Finally, a large and continuous Würm distribution is highly likely south of the Fennoscandian glaciers in Central Europe for C. palaeno, which might indicate extended peatland areas during Würm glacial. With all the patterns found in this study, the understanding of glacial persistence of forest, respectively forest-like structures and peatlands during Würm or even Riss glacial in Europe could be advanced. The congruencies among the analysed woodland and bog species illustrate the importance and location of extra-Mediterranean refugia for European mountain forests and the glacial presence of Central European peatlands. Thus, already postulated theories could be supported and further pieces of the overall puzzle could be added. The varieties of the different survival centres once more clarified that further phylogeographic studies on mountain forest of different habitat requirements and especially peatland species have to be implemented to get a clearer picture of the glacial history of these habitats.