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Institute
Stress has been considered one of the most relevant factors promoting aggressive behavior. Animal and human pharmacological studies revealed the stress hormones corticosterone in rodents and cortisol in humans to constitute a particularly important neuroendocrine determinate in facilitating aggression and beyond that, assumedly in its continuation and escalation. Moreover, cortisol-induced alterations of social information processing, as well as of cognitive control processes, have been hypothesized as possible influencing factors in the stress-aggression link. So far, the immediate impact of a preceding stressor and thereby stress-induced rise of cortisol on aggressive behavior as well as higher-order cognitive control processes and social information processing in this context have gone mostly unheeded. The present thesis aimed to extend the hitherto findings of stress and aggression in this regard. For this purpose two psychophysiological studies with healthy adults were carried out, both using the socially evaluated-cold pressor test as an acute stress induction. Additionally to behavioral data and subjective reports, event related potentials were measured and acute levels of salivary cortisol were collected on the basis of which stressed participants were divided into cortisol-responders and "nonresponders. Study 1 examined the impact of acute stress-induced cortisol increase on inhibitory control and its neural correlates. 41 male participants were randomly assigned to the stress procedure or to a non-stressful control condition. Beforehand and afterwards, participants performed a Go Nogo task with visual letters to measure response inhibition. The effect of acute stress-induced cortisol increase on covert and overt aggressive behavior and on the processing of provoking stimuli within the aggressive encounter was investigated in study 2. Moreover, this experiment examined the combined impact of stress and aggression on ensuing affective information processing. 71 male and female participants were either exposed to the stress or to the control condition. Following this, half of each group received high or low levels of provocation during the Taylor Aggression Paradigm. At the end of the experiment, a passive viewing paradigm with affective pictures depicting positive, negative, or aggressive scenes with either humans or objects was realized. The results revealed that men were not affected by a stress-induced rise in cortisol on a behavioral level, showing neither impaired response inhibition nor enhanced aggressive behavior. In contrast, women showed enhanced overt and covert aggressive behavior under a surge of endogenous cortisol, confirming previous results, albeit only in case of high provocation and only up to the level of the control group. Unlike this rather moderate impact on behavior, cortisol showed a distinct impact on neural correlates of information processing throughout inhibitory control, aggression-eliciting stimuli, and emotional pictures for both men and women. At this, stress-induced increase of cortisol resulted in enhanced N2 amplitudes to Go stimuli, whereas P2 amplitudes to both and N2 to Nogo amplitudes retained unchanged, indicating an overcorrection and caution of the response activation in favor of successful inhibitory control. The processing of aggression-eliciting stimuli during the aggressive encounter was complexly altered by stress differently for women and men. Under increased cortisol levels, the frontal or parietal P3 amplitude patterns were either diminished or reversed in the case of high provocation compared to the control group and to cortisol-nonresponders, indicating a desensitization towards aggression-eliciting stimuli in males, but a more elaborate processing of those in women. Moreover, stress-induced cortisol and provocation jointly altered subsequent affective information processing at early as well as later stages of the information processing stream. Again, increased levels of cortisol led opposite directed amplitudes in the case of high provocation relative to the control group and cortisol-nonresponders, with enhanced N2 amplitudes in men and reduced P3 and LPP amplitudes in men and women for all affective pictures, suggesting initially enhanced emotional reactivity in men, but ensuing reduced motivational attention and enhanced emotion regulation in both, men and women. As a result, these present findings confirm the relevance of HPA activity in the elicitation and persistence of human aggressive behavior. Moreover, they reveal the significance of compensatory and emotion regulatory strategies and mechanisms in response to stress and provocation, indorsing the relevance of social information and cognitive control processes. Still, more research is needed to clarify the conditions which lead to the facilitation of aggression and by which compensatory mechanisms this is prevented.
A sustainable development of forests and their ecosystem services requires the monitoring of the forests" state and changes as well as the prediction of their future development. To achieve the latter, eco-physiological forest growth models are usually applied. These models require calibration and validation with forestry reference data. This data includes forest structural parameters such as tree height or stem diameter which are easy to measure and can be used to estimate the core model parameters, i.e. the tree- biomass pools. The methods traditionally applied to derive the structural parameters are mainly manual and time-consuming. Hence, the in situ data acquisition is inefficient and limited in its ability to capture the vertical and horizontal variability in stand structure. Ground-based remote sensing bears the potential to overcome the limitations of the traditional methods. As they can be automated, ground-based remote sensing methods allow a much more efficient data acquisition and a larger spatial coverage. They are also able to capture forest structure in its three dimensions. Nevertheless, at present further research is required, in particular with respect to the practical integration of ground-based remote sensing data into forest growth models as well as regarding factors influencing the structural parameter retrieval from this data. Therefore, the goal of this PhD thesis was to investigate the influencing factors of two ground-based remote sensing methods (terrestrial laser scanning and hemispherical photography), which have not or only scarcely been studied to date. In addition, the use of forest structural parameters derived from these methods for the calibration of a forest growth model was assessed. Both goals were achieved. The results of this thesis could contribute significantly to a comprehensive assessment of ground-based remote sensing and its potential to derive the forest structural parameters. However, the use of these methods to calibrate forest growth models proved to be limited. An optimized data sampling design is expected to eliminate the major limitations, though. Furthermore, the combination of ground-based, airborne, and satellite remote sensing sensors was suggested to provide an optimized framework for the general integration of remotely sensed data into forest growth models. This combination of remote sensing observations at different scales will contribute greatly to a modern forest management with the purpose of warranting a sustainable forest development even under growing economic and ecological pressures.
Attitudes are "the most distinctive and indispensable concept in contemporary social psychology" (Allport, 1935, p. 798). This outstanding position of the attitude concept in social cognitive research is not only reflected in the innumerous studies focusing on this concept but also in the huge number of theoretical approaches that have been put forth since then. Yet, it is still an open question, what attitudes actually are. That is, the question of how attitude objects are represented in memory cannot be unequivocally answered until now (e.g., Barsalou, 1999; Gawronski, 2007; Pratkanis, 1989, Chapter 4). In particular, researchers strongly differ with respect to their assumptions on the content, format and structural nature of attitude representations (Ferguson & Fukukura, 2012). This prevailing uncertainty on what actually constitutes our likes and dislikes is strongly dovetailed with the question of which processes result in the formation of these representations. In recent years, this issue has mainly been addressed in evaluative conditioning research (EC). In a standard EC-paradigm a neutral stimulus (conditioned stimulus, CS) is repeatedly paired with an affective stimulus (unconditioned stimulus, US). The pairing of stimuli then typically results in changes in the evaluation of the CS corresponding to the evaluative response of the US (De Houwer, Baeyens, & Field, 2005). This experimental approach on the formation of attitudes has primarily been concerned with the question of how the representations underlying our attitudes are formed. However, which processes operate on the formation of such an attitude representation is not yet understood (Jones, Olson, & Fazio, 2010; Walther, Nagengast, & Trasselli, 2005). Indeed, there are several ideas on how CS-US pairs might be encoded in memory. Notwithstanding the importance of these theoretical ideas, looking at the existing empirical work within the research area of EC (for reviews see Hofmann, De Houwer, Perugini, Baeyens, & Crombez, 2010; De Houwer, Thomas, & Baeyens, 2001) leaves one with the impression that scientists have skipped the basic processes. Basic processes hereby especially refer to the attentional processes being involved in the encoding of CSs and USs as well as the relation between them. Against the background of this huge gap in current research on attitude formation, the focus of this thesis will be to highlight the contribution of selective attention processes to a better understanding of the representation underlying our likes and dislikes. In particular, the present thesis considers the role of selective attention processes for the solution of the representation issue from three different perspectives. Before illustrating these different perspectives, Chapter 1 is meant to envision the omnipresence of the representation problem in current theoretical as well as empirical work on evaluative conditioning. Likewise, it emphasizes the critical role of selective attention processes for the representation question in classical conditioning and how this knowledge might be used to put forth the uniqueness of evaluative conditioning as compared to classical conditioning. Chapter 2 then considers the differential influence of attentional resources and goal-directed attention on attitude learning. The primary objective of the presented experiment was thereby to investigate whether attentional resources and goal-directed attention exert their influence on EC via changes in the encoding of CS-US relations in memory (i.e., contingency memory). Taking the findings from this experiment into account, Chapter 3 focuses on the selective processing of the US relative to the CS. In particular, the two experiments presented in this chapter were meant to explore the moderating influence of the selective processing of the US in its relation to the CS on EC. In Chapter 4 the important role of the encoding of the US in relation to the CS, as outlined in Chapter 3, is illuminated in the context of different retrieval processes. Against the background of the findings from the two presented experiments, the interplay between the encoding of CS-US contingencies and the moderation of EC via different retrieval processes will be discussed. Finally, a general discussion of the findings, their theoretical implications and future research lines will be outlined in Chapter 5.
Evaluative conditioning (EC) refers to changes in liking that are due to the pairing of stimuli, and is one of the effects studied in order to understand the processes of attitude formation. Initially, EC had been conceived of as driven by processes that are unique to the formation of attitudes, and that occur independent of whether or not individuals engage in conscious and effortful propositional processes. However, propositional processes have gained considerable popularity as an explanatory concept for the boundary conditions observed in EC studies, with some authors going as far as to suggest that the evidence implies that EC is driven primarily by propositional processes. In this monograph I present research which questions the validity of this claim, and I discuss theoretical challenges and avenues for future EC research.
Mankind has dramatically influenced the nitrogen (N) fluxes between soil, vegetation, water and atmosphere " the global N cycle. Increasing intensification of agricultural land use, caused by the growing demand for agricultural products, has had major impacts on ecosystems worldwide. Particularly nitrogenous gases such as ammonia (NH3) have increased mainly due to industrial livestock farming. Countries with high N deposition rates require a variety of deposition measurements and effective N monitoring networks to assess N loads. Due to high costs, current "conventional"-deposition measurement stations are not widespread and therefore provide only a patchy picture of the real extent of the prevailing N deposition status over large areas. One tool that allows quantification of the exposure and the effects of atmospheric N impacts on an ecosystem is the use of bioindicators. Due to their specific physiology and ecology, especially lichens and mosses are suitable to reflect the atmospheric N input at ecosystem level. The present doctoral project began by investigating the general ability of epiphytic lichens to qualify and quantify N deposition by analysing both lichens and total N and δ15N along a gradient of different N emission sources and severity. The results showed that this was a viable monitoring method, and a grid-based monitoring system with nitrophytic lichens was set up in the western part of Germany. Finally, a critical appraisal of three different monitoring techniques (lichens, mosses and tree bark) was carried out to compare them with national relevant N deposition assessment programmes. In total 1057 lichen samples, 348 tree bark samples, 153 moss samples and 24 deposition water samples, were analysed in this dissertation at different investigation scales in Germany.The study identified species-specific ability and tolerance of various epiphytic lichens to accumulate N. Samples of tree bark were also collected and N accumulation ability was detected in connection with the increased intensity of agriculture, and according to the presence of reduced N compounds (NHx) in the atmosphere. Nitrophytic lichens (Xanthoria parietina, Physcia spp.) have the strongest correlations with high agriculture-related N deposition. In addition, the main N sources were revealed with the help of δ15N values along a gradient of altitude and areas affected by different types of land use (NH3 density classes, livestock units and various deposition types). Furthermore, in the first nationwide survey of Germany to compare lichens, mosses and tree bark samples as biomonitors for N deposition, it was revealed that lichens are clearly the most meaningful monitor organisms in highly N affected regions. Additionally, the study shows that dealing with different biomonitors is a difficult task due to their variety of N responses. The specific receptor surfaces of the indicators and therefore their different strategies of N uptake are responsible for the tissue N concentration of each organism group. It was also shown that the δ15N values depend on their N origin and the specific N transformations in each organism system, so that a direct comparison between atmosphere and ecosystems is not possible.In conclusion, biomonitors, and especially epiphytic lichens may serve as possible alternatives to get a spatially representative picture of the N deposition conditions. Furthermore, bioindication with lichens is a cost-efficient alternative to physico-chemical measurements to comprehensively assess different prevailing N doses and sources of N pools on a regional scale. They can at least support on-site deposition instruments by qualification and quantification of N deposition.
High-resolution projections of the future climate are required to assess climate change realistically at a regional scale. This is in particular important for climate change impact studies since global projections are much too coarse to represent local conditions adequately. A major concern is thereby the change of extreme values in a warming climate due to their severe impact on the natural environment, socio-economical systems and the human health. Regional climate models (RCMs) are, however, able to reproduce much of those local features. Current horizontal resolutions are about 18-25km, which is still too coarse to directly resolve small-scale processes such as deep-convection. For this reason, projections of a possible future climate were simulated in this study with the regional climate model COSMO-CLM at horizontal resolutions of 4.5km and 1.3km for the region of Saarland-Lorraine-Luxemburg and Rhineland-Palatinate for the first time. At a horizontal scale of about 1km deep-convection is treated explicitly, which is expected to improve particularly the simulation of convective summer precipitation and a better resolved orography is expected to improve near surface fields such as 2m temperature. These simulations were performed as 10-year long time-slice experiments for the present climate (1991"2000), the near future (2041"2050) and the end of the century (2091"2100). The climate change signals of the annual and seasonal means and the change of extremes are analysed with respect to precipitation and 2m temperature and a possible added value due to the increased resolution is investigated. To assess changes in extremes, extreme indices have been applied and 10- and 20-year return levels were estimated by "peak-over-threshold" models. Since it is generally known that model output of RCMs should not directly be used for climate change impact studies, the precipitation and temperature fields were bias-corrected with several quantile-matching methods. Among them is a new developed parametric method which includes an extension for extreme values and is hence expected to improve the correction. In addition, the impact of the bias-correction on the climate change signals and on the extreme value statistics was investigated. The results reveal a significant warming of the annual mean by about +1.7 -°C until 2041"2050 and +3.7 -°C until 2091"2100, but considerably stronger signals of up to +5 -°C in summer in the Rhine Valley. Furthermore, the daily variability increases by about +0.8 -°C in summer but decreases by about -0.8 -°C in winter. Consequently, hot extremes increase moderately until the mid of the century but strongly thereafter, in particular in the Rhine Valley. Cold extremes warm continuously in the complete domain in the next 100 years but strongest in mountainous areas. The change signals with regard to annual precipitation are of the order -±10% but not significant. Significant, however, are a predicted increase of +32% of the seasonal precipitation in autumn until 2041"2050 and a decrease of -28% in summer until 2091-2100. No significant changes were found for days with intensities > 20 mm/day, but the results indicate that extremes with return periods ≤2 years increase as well as the frequency and duration of dry periods. The bias-corrections amplified positive signals but dampened negative signals and considerably reduced the power of detection. Moreover, absolute values and frequencies of extremes were altered by the correction but change signals remained approximately constant. The new method outperformed other parametric methods, in particular with regard to extreme value correction and related extreme indices and return levels. Although the bias correction removed systematic errors, it should be treated as an additional layer of uncertainty in climate change studies. Finally, the increased resolution of 1.3km improved predominantly the representation of temperature fields and extremes in terms of spatial heterogeneity. The benefits for summer precipitation were not as clear due to a severe dry-bias in summer, but it could be shown that in principle the onset and intensity of convection improves. This work demonstrates that climate change will have severe impacts in this investigation area and that in particular extremes may change considerably. An increased resolution provides thereby an added value to the results. These findings encourage further investigations, for other variables as for example near-surface wind, which will be more feasible with growing computing resources. These analyses should, however, be repeated with longer time series, different RCMs and anthropogenic scenarios to determine the robustness and uncertainty of these results more extensively.
Veterinary antibiotics are released to arable agricultural soil together with manure, including nutrients, organic matter, and microorganisms. Previously, the effects of antibiotic-contaminated manure on soil microbial community activity, function, structure, and resistance have been reported under controlled experimental conditions. This thesis further evaluated the antimicrobial effects as influenced by different manure compositions, soil microhabitats and moisture regimes, plants, and different distances to roots. Microbial community responses were determined by phenotypic phospholipid fatty acid (PLFA) and genotypic 16S rRNA gene fragment analyses. (Chapter 3) demonstrates that medication of pigs with difloxacin (DIF) and sulfadiazine (SDZ) alters the molecular-chemical pattern of slurries, confounding the detection of a consistent antibiotic effect in bulk and respective rhizosphere soil. This was evaluated in a 63-day mesocosm experiment considering typical agricultural manure applications to maize planted soil. Fecal bacteria were detected even 14 days after manure amendment. Manure of DIF- and SDZ-medicated pigs clearly affected the microbial community in mesocosm bulk and rhizosphere soil, temporarily matching antibiotic effects reported in previous studies. (Chapter 4) discusses the influences of different soil microhabitats on antibiotic fate and the effects on soil microflora. Total extractable SDZ was more than two-fold larger in earthworm burrows and soil macroaggregate surfaces compared to bulk soil or the interior fraction of aggregates. Furthermore, soil microbial communities were affected by a combination of soil microhabitat and treatment, which was reflected by different structural and functional community responses to SDZ in laboratory and under field conditions. (Chapter 5) evaluates if SDZ effects on microbial communities are more pronounced in soils which undergo periodic changes in soil moisture by drying-rewetting dynamics compared to soils without such moisture fluctuations. This was tested in a 49-day climate chamber soil pot experiment grown with grass. Manure-amended pots without or with SDZ contamination were incubated under a dynamic moisture regime with repeated drying and rewetting changes of more than twenty percent maximum water holding capacity compared to the control moisture regime. The microbial biomass, but less pronouncedly the community structure, showed an increased responsiveness to the combined stress of SDZ and dynamic moisture changes in the laboratory. Similar responses were documented under field conditions. (Chapter 6) indicated adverse effects of SDZ on root geotropism, number of lateral roots, and water uptake by plants in a 40-day greenhouse experiment with willow and maize grown in soil with environmentally relevant and worst-case antibiotic contamination. (Chapter 7) showed that the associated microbial community responded to a combination of plant species, distance to the root, and antibiotic spiking concentration. In highly antibiotic-contaminated soils, the structural and functional responses of the microbial community were dominated by indirect antibiotic effects on plants and roots.
Fast and Slow Effects of Cortisol on Several Functions of the Central Nervous System in Humans
(2014)
Cortisol is one of the key substances released during stress to restore homeostasis. Our knowledge of the impact of this glucocorticoid on cognition and behavior in humans is, however, still limited. Two modes of action of cortisol are known, a rapid, nongenomic and a slow, genomic mode. Both mechanisms appear to be involved in mediating the various effects of stress on cognition. Here, three experiments are presented that investigated fast and slow effects of cortisol on several functions of the human brain. The first experiment investigated the interaction between insulin and slow, genomic cortisol effects on resting regional cerebral blood flow (rCBF) in 48 young men. A bilateral, locally distinct increase in rCBF in the insular cortex was observed 37 to 58 minutes after intranasal insulin admission. Cortisol did not influence rCBF, neither alone nor in interaction with insulin. This finding suggests that cortisol does not influence resting cerebral blood flow within a genomic timeframe. The second experiment examined fast cortisol effects on memory retrieval. 40 participants (20 of them female) learned associations between neutral male faces and social descriptions and were tested for recall one week later. Cortisol administered intravenously 8 minutes before retrieval influenced recall performance in an inverted U-shaped dose-response relationship. This study demonstrates a rapid, presumably nongenomic cortisol effect on memory retrieval in humans. The third experiment studied rapid cortisol effects on early multisensory integration. 24 male participants were tested twice in a focused cross-modal choice reaction time paradigm, once after cortisol and once after placebo infusion. Cortisol acutely enhanced the integration of visual targets and startling auditory distractors, when both stimuli appeared in the same sensory hemi-field. The rapidity of effect onset strongly suggests that cortisol changes multisensory integration by a nongenomic mechanism. The work presented in this thesis highlights the essential role of cortisol as a fast acting agent during the stress response. Both the second and the third experiment provide new evidence of nongenomic cortisol effects on human cognition and behavior. Future studies should continue to investigate the impact of rapid cortisol effects on the functioning of the human brain.
In splitting theory of locally convex spaces we investigate evaluable characterizations of the pairs (E, X) of locally convex spaces such that each exact sequence 0 -> X -> G -> E -> 0 of locally convex spaces splits, i.e. either X -> G has a continuous linear left inverse or G -> E has a continuous linear right inverse. In the thesis at hand we deal with splitting of short exact sequences of so-called PLH spaces, which are defined as projective limits of strongly reduced spectra of strong duals of Fréchet-Hilbert spaces. This class of locally convex spaces contains most of the spaces of interest for application in the theory of partial differential operators as the space of Schwartz distributions , the space of real analytic functions and various spaces of ultradifferentiable functions and ultradistributions. It also contains non-Schwartz spaces as B(2,k,loc)(Ω) and spaces of smooth and square integrable functions that are not covered by the current theory for PLS spaces. We prove a complete characterizations of the above problem in the case of X being a PLH space and E either being a Fréchet-Hilbert space or a strong dual of one by conditions of type (T ). To this end, we establish the full homological toolbox of Yoneda Ext functors in exact categories for the category of PLH spaces including the long exact sequence, which in particular involves a thorough discussion of the proper concept of exactness. Furthermore, we exhibit the connection to the parameter dependence problem via the Hilbert tensor product for hilbertizable locally convex spaces. We show that the Hilbert tensor product of two PLH spaces is again a PLH space which in particular proves the positive answer to Grothendieck- problème des topologies. In addition to that we give a complete characterization of the vanishing of the first derivative of the functor proj for tensorized PLH spectra if one of the PLH spaces E and X meets some nuclearity assumptions. To apply our results to concrete cases we establish sufficient conditions of (DN)-(Ω) type and apply them to the parameter dependence problem for partial differential operators with constant coefficients on B(2,k,loc)(Ω) spaces as well as to the smooth and square integrable parameter dependence problem. Concluding we give a complete solution of all the problems under consideration for PLH spaces of Köthe type.
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.