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This thesis draws a comprehensive picture about the radiation and diversification of truncatelloidean gastropods across the south pacific. It covers three more specifc studies focussing on the Truncelloideans from Fiji, Vanuatu and New Caledonia, respectively. And a conclusive analysis that combines the results of the three more specific studies and enhances them using species from the Austral Islands, Lord Howe Island, the Indonesian island Sulawesi as well as several species from New Zealand and Australia. Molecular phylogenies were calculated using four nuclear gene fragments (ITS2; 18S rRNA; 28S rRNA and Histone 3) besides the mitochondrial COI and 16S rRNA. Further molecuular data was used to calculate dated phylogenies, perform ancestral range reconstructions and develop a modified molecular barcoding approach.
The need for the diversification of utilised species has emerged in the present aquaculture
production environment. Shifts in consumer interest, climate change-induced temperature
increases, and major fish disease outbreaks have put a strain on this industry. In this context,
the pikeperch (Sander lucioperca) has become a new target species for aquaculture in Central
Europe. This new aquaculture focus species exhibits high numbers of offspring, fast growth,
and high consumer acceptance. It can also effectively deal with higher temperatures and turbid
water. However, the rate of successful rearing is still low, as various developmental
transformations and environmental effects commonly lead to high mortality rates during the
early ontogenetic stages. The aim of this doctoral project was thus to obtain insight into
embryonic to larval developmental changes during pikeperch ontogeny. Specifically, the times
of change that influence survival were of focus. Based on the available literature, particular
attention was paid to general growth patterns and the connected developmental changes, the
determination of myogenesis gene marker expression changes, and the support of animal
welfare efforts for pikeperch rearing procedures. To achieve the aims of the study, a methodical
setup consisting of morphometric and developmental observations was combined with
transcriptome gene marker analysis for the different ontogenetic stages.
Three developmental phases were differentiated during the embryo-larval transition. Each of
these possessed distinct growth patterns with different growth rates. The intermediate
threshold phase showed internal organ development that focused on digestive, neuronal, and
heart tissues. Three activity phases of myogenesis were determined: during early embryonic
development, before hatching, and after hatching during the larval stages. Therefore, muscle
development seemed to be regulated to balance energy expenditures. Additionally, two
coinciding skeletogenic phases were found. Furthermore, a cell line from whole embryos was
developed to support the replacement of animals in future experimental setups. A software
system for video analyses was developed to support rearing procedures in aquaculture
facilities. This prototype can be used to automate the counting of specimens and thus allows
for faster responses to increasing mortalities. Based on the results of this thesis project, further
insights into the early development of pikeperches were obtained. This will facilitate the design
and adaptation of raising and husbandry protocols, which can help to further establish
pikeperch as an aquaculture species and support its application in modern recirculatory
systems.
Having been regarded as wastelands until quite recently, wetlands are increasingly acknowledged as ecosystems of high biodiversity. Wetland restoration projects are often accompanied by the implementation of specific species management programs. Naturally, for effective management measures, profound knowledge of the target speciesʼ ecological requirements is obligatory, including habitat selection, feeding ecology as well as spatial behaviour such as movements within and between patches of suitable habitat. Yet, big knowledge gaps exist for many marshland birds which is particularly true for highly secretive species such as rails and crakes. Considered as the least known among the Palaearctic breeding birds, most information about the Baillon's Crake Zapornia pusilla is only anecdotic, resulting in strong uncertainties with regard to the species' distribution, population sizes, status, migratory behaviour as well as ecological requirements. This can be mainly attributed to the species' skulking behaviour and its seemingly highly erratic occurrence. Baillon's Crakes in the Western Palaearctic and Palaeotropics are referred to as the subspecies Z. p. intermedia. While European breeding birds are assumed to winter in sub-Saharan wetlands, African populations are considered rather to be itinerant with local movements induced by seasonal or anthropogenic habitat changes. However, for both migratory movements, major directions or routes are unknown. The discovery of a large number of Baillon's Crakes presumably wintering in the floodplains of the Parc National des Oiseaux du Djoudj (PNOD), situated in the Senegal River Delta, WAfrica, initiated this thesis. The main aim of the study was, firstly, to clarify the status and size of this population and assess its connectivity to European breeding population(s). Secondly, in order to improve the knowledge about the species' ecological requirements as a basis for the National Parks conservation management, habitat selection, spatial behaviour as well as dietary selectivity were investigated. The major part of the fieldwork was performed in PNOD in the course of the dry season during periods of 1.5 - 2.5 months from December - March 2009, 2010 and 2013. Baillon's Crakes were mainly caught with cage traps, ringed and common measurements were taken, including moult status. Skin tissue as well as one rectrice was sampled for DNA and stable isotope analyses. This was also done for Baillon's Crakes caught in European breeding grounds in Germany, Montenegro and Southern Spain. For dietary analyses, faecal samples were collected in PNOD in winter 2009/2010. Furthermore, some individuals were equipped with radio-transmitters to determine home range size and habitat selection. For the identification of the most relevant habitat parameters both on a population as well as on the individuals' level, we used a vegetation map based on satellite imagery covering the entire Djoudj area as well as maps generated on the basis of aerial photographs taken at two study sites.
Presumably every organism on earth is involved in at least one mutualistic interaction with one or several other species. To interact with each other, the species need traits that provide benefits to the partner species. Surprisingly, the function of traits for the stabilization of mutualisms has rarely been investigated, despite of a general lack of knowledge how mutualisms are maintained. The aim of this work was to find functional traits, which stabilize the mutualism between a bat species and a carnivorous pitcher plant in Northern Borneo. Kerivoula hardwickii is the only bat species known to roost in pitcher-shaped trapping organs of Palaeotropical pitcher plants (Nepenthes). These bats fertilize the pitcher plant Nepenthes hemsleyana with their nutritious nitrogen-rich faeces while roosting inside the pitchers. The plants have outsourced capture and digestion of arthropod prey to the bats on which they strongly rely for nutrient acquisition. The bats in contrast are less dependent on their mutualism partner as they also roost in pitchers of two further Nepenthes species as well as in developing furled leaves of various plant species in the order Zingiberales. In earlier studies, we found that N. hemsleyana outcompetes alternative roosts by providing high-quality roosts for the bats. However, which traits exactly stabilize the mutualism between K. hardwickii and N. hemsleyana was still unclear. I found that both the bats and the pitcher plants show traits, which have the potential to stabilize their interaction. On the level of morphological traits, I found that the pitchers have a low fluid level and a particular shape that provide just enough roosting space for one individual of the solitary K. hardwickii, a mother with juvenile or a mating couple. The bats have enlarged thumb and foot pads that enable them to cling to the smooth surfaces of their roosts without using their claws. This avoids damage to the sensitive N. hemsleyana pitchers. On the level of communicational traits, again N. hemsleyana acquired morphological structures that act as effective ultrasound-reflectors, which guide the echo-orientating bats to the opening of the pitchers and help the bats to identify their mutualism partner. The bats’ calls on the other hand are characterized by extraordinary high starting frequencies and broad bandwidths, which enable K. hardwickii to easily locate pitchers of N. hemsleyana and other Nepenthes species in their dense habitats. Finally, on the level of behavioural traits the bats often but not always prefer their mutualism partner to other roosts when they can select roosts in their natural environment or in behavioural experiments. The reason for this behaviour seems to be a combination of 1) N. hemsleyana’s superior quality compared to alternative roosts and 2) different roosting traditions of the bats. In conclusion, the mutualism between bats and pitcher plants is asymmetric as N. hemsleyana is more dependent on K. hardwickii than vice versa. For the plants bat faeces present their most important nutrient source. In contrast, K. hardwickii can select between alternative roosting plants. This asymmetric dependency is reflected in the specifity and function of the traits that stabilize the mutualism in each of the two involved species. Especially on the morphological level, N. hemsleyana seems to have evolved several traits that perfectly fit to K. hardwickii. In contrast, the bats’ traits more generally facilitate their roosting in funnel-shaped plant structures and their occurrence in cluttered habitats. Thus, they are probably exaptations (i.e. traits that evolved for another reason) that are nevertheless functional and stabilize the mutualism with N. hemsleyana. This plant‘s superior roost quality is likely a consequence of the competition with alternative roosting plants and is a pre-requisite for the bats to prefer N. hemsleyana. Moreover, my study confirms earlier findings that asymmetric dependencies support the stabilization of mutualistic interactions. Finally, my work indicates that the specifity of functional traits can be used as a measure to determine mutual dependencies of mutualistic partners.
Bats belong to the most gregarious and diverse mammals with highly complex social behaviors. Despite extensive research on their ecology and social behavior in some bat species, gained insights are restricted to only few of the more than 1300 species. In the recent past, bats have also become a central topic of a different branch of research: Since the 1990s bats came to the fore of virologists and immunologists due to the bats’ apparent importance as reservoir hosts and vectors of several (mostly tropical) diseases. While this research is focused mainly on emerging infectious diseases linked to bats, and their zoonotic potential, little has been invested regarding the link between disease transmission and bat social systems.
In my work, I aim at filling this gap by merging automated daily roosting observations, social network analysis, and a virological screening in Natterer’s bats (Myotis nattereri). In a collaborative approach, my co-workers and I analyzed the social structure of individually marked Natterer’s bats, their astrovirus detection rate and transmission pathways within their colony, as well as roosting interactions between different co-occurring con- and heterospecific bat colonies.
We discovered Natterer’s bats to display a very divergent social network structure that contradicts the findings of previous studies on large fission-fusion groups. Contrary to the modular social network structure found in e.g. primates or other bats species, the social network of Natterer’s bats consists of only one highly interconnected community. Moreover, although the close proximity between bat hosts in the colony should strongly promote direct transmission, we found indications that astrovirus infections follow at least partly an indirect transmission pathway via contaminated roost use. Lastly, our results prove that co-occurring con- and heterospecific bat colonies, e.g. as in this study Natterer’s bats, brown long-eared bats and Bechstein’s bats, can influence each other in their roost use by avoiding conspecific roosts and by being attracted towards those of heterospecifics. This holds implication for the transmission of parasites and pathogens within and between different colonies with opportunities for spillovers. To conclude, this multidisciplinary study led to valuable insights in the hitherto hidden mechanisms within and among bat colonies.
Dwarf spiders (Linyphiidae, Erigoninae) are especially suitable for sexual selection research as many of them exhibit sexual dimorphism, with males possessing modified prosomata. In those species that have been investigated in detail the modified structures are equipped with a glandular tissue that produces secretions, which the females contact and take up during courtship/copulation. The time of secretion release, and refilling of the reservoirs was analysed on an ultrastructural level in male Oedothorax retusus. The results suggest that the main function of the secretions is gustatorial courtship and not the emission of volatile pheromones for mate attraction. Mating decisions and reproductive success are influenced by secondary sexual traits that evolved under sexual selection. However, an individual´s nutritional status is also important for mate choice. Since spiders are regularly exposed to limited prey availability, adult feeding status can be considered an important component of spider mating behaviour. In order to test for the effects of dietary restriction, females of the closely related species O. retusus and O. apicatus were subject to a short period of food shortage. The effects of low- (LD) vs. high-diet (HD) treatment on courtship, mating probability and behaviour, and reproduction were analysed. We found that short phases of diet restriction as adults have a high impact on copulation and reproduction in the two dwarf spider species. Whenever females mate with more than one male, and sperm is stored prior to fertilization, males may suffer from sperm competition. Mating plugs that block the female genital openings after mating are a male strategy to avoid sperm competition. Although mating plugs occur in many species, their function and origin has hardly been investigated. O. retusus males transfer amorphous material onto the female genitalia during mating. We investigated the location of plug production using x-ray microtomography (μCT) as well as light and transmission electron microscopy (TEM). Furthermore, we asked whether males are limited in the production of the amorphous plug material in successive matings. The plug material is produced in a gland inside the male pedipalp and stored close to the blind end of the sperm storage compartment. The size of the first plug a male produced significantly influenced the size of subsequent plugs. Obviously, males do not possess unlimited amounts of plug material in a certain period, which may severely limit their ability to secure paternity through subsequent mating plugs. Even though mating plugs seem to be an obvious means to secure paternity, their potential in securing paternity depends on their mechanical efficacy and persistence. Consequently, the influence of the size of the plug material (mating duration as a proxy) and the age of the mating plug (time interval between successive copulations) on its efficacy was investigated. Small and fresh plugs were least effective, whereas large plugs were highly effective. We were able to show that mating plugs in O. retusus are a powerful mechanical safeguard whose efficacy varies with plug size and age. Genitalia in animals with internal fertilization are complex, species-specific, and underlie rapid evolution. In spiders, male and female genitalia are paired, and have to interact during mating, which results in an even higher complexity. Pedipalps (transformed pair of legs) in male spiders are used as secondary sperm transfer organs that are not directly connected to the gonads. Due to the high complexity of male pedipalps, it has been taken for granted that pedipalps are side specific and cannot be used flexibly into either female copulatory opening. We investigated potential flexible pedipalp use in O. retusus. Our findings demonstrate a flexible insertion mode in a dwarf spider with complex pedipalps but relatively simple female genitalia. Our findings corroborate sexual selection as the selective regime for the evolution of complex and diverse genitalia. The results of this thesis show how complex sexual selection acts in the dwarf spiders O. retusus and O. apicatus. It shapes the evolution of male and female genitalia, affects mate choice (pre- and postcopulatory), mating behaviour, and influences mating success and reproduction. All these factors and traits affect an individual´s evolutionary fitness, and their interactions help to understand how sexual selection acts.
Relative importance of plastic and genetic responses to weather conditions in long-lived bats
(2022)
In the light of the accelerating pace of environmental change, it is imperative to understand how populations and species can adapt to altered environmental conditions. This is a crucial step in predicting current and future population persistence and limits thereof. Genetic adaption and phenotypic plasticity are two main mechanisms that can mediate the process of adaptation and are of particular importance for non-dispersing species. While phenotypic plasticity may enable individuals to cope with short term environmental changes, genetic adaptation will often be required for populations to survive in situ over longer time spans. However, a rapid genetic response is expected particularly in species with fast life histories or large population sizes, leaving species with slow life histories potentially at higher extinction risk. The Bechstein’s bat (Myotis bechsteinii) is a mammal of 10 g weight that - despite its small size - is characterized by a slow life history, with low reproductive output and long lifespan, and is already considered to be of high conservation concern. Past work demonstrated body size to be a highly fitness-relevant trait in Bechstein’s bats. Body size is further known to be a pivotal trait shaping the pace of life histories in numerous species. Simultaneously, many studies reported noteworthy changes in body size as a response to shifting environments across different taxa. This suggested a potential for high plasticity in this trait in Bechstein’s bats as well; however, changes in body size could have vital impacts on demographic rates.
Therefore, this dissertation investigated the following questions: firstly, what shapes the fundamental development of body size in M. bechsteinii, and, specifically, is there an impact of weather conditions on body size? If so, in what form and magnitude? Secondly, how does body size subsequently influence the pace of life in females? What is the cost of a faster or slower pace of life, and how does fitness compare across individuals with slow and fast life histories? And finally, to what extent can changes in body size be attributed to either phenotypic plasticity or genetic adaptation? What is the evolutionary potential of body size in the populations? And, consequently, what implications can we draw regarding population persistence of these colonies?
To answer these questions, we analyzed a long-term dataset of over two decades collected from four wild Bechstein’s bat colonies. We used individual-based data on survival, reproduction and body size, built multi-generational pedigrees, and combined everything with meteorological data. In Manuscript 1 we found that, in contrast to the declining body size observed in many species, body size in Bechstein’s bats increased significantly over the last decades. We demonstrated that ambient temperature was linked to the development of body size and identified a sensitive time period in the prenatal growth phase, in which body size was most susceptible to the impact of temperature. We established that warmer summers resulted in larger bats, but that these large bats had higher mortality risks throughout their lives. Manuscript 2 then revealed the influence of body size on the pace of life in Bechstein’s bats and demonstrated high plasticity in intraspecific life history strategies. Large females were characterized by a faster pace of life and shorter lifespans, but surprisingly, lifetime reproductive success remained remarkably stable across individuals with different body sizes. The acceleration of their pace of life means that larger females compensated for their reduced longevity by an earlier reproduction and higher fecundity to reach similar overall fitness. Ultimately, differences in body size resulted in changes in population growth rate via the impact of size on generation times. Results of Manuscript 3 were then able to clarify the extent to which changes in body size were founded on either phenotypic plasticity or genetic adaptation. We demonstrated a particularly low heritability in hot summers, indicating that variance in body size was mostly driven by phenotypic plasticity, with few genetic constraints. During cold summers, behavioural adaptations by reproducing bats seem to be able to mitigate negative effects of cold temperatures. These behaviours, such as social aggregation or preference for warm roosts, are, however, essentially irrelevant in hot environments. In addition, a low evolvability of forearm length points to a low capacity to respond to selection pressures associated with the trait.
We can conclude that body size in M. bechsteinii has increased over the last two decades as a response to global warming and is only slightly constrained by its genetic underpinnings. We can further demonstrate a direct link between body size and the pace of life histories in the Bechstein’s bat populations and how changes in body size impact demographic rates via this linkage. In the context of climate change and hotter summers, our findings consequently suggest that body size will likely increase further if warm summers continue to become more frequent. Whether this plastic response of body size proves to be adaptive in the long term, however, remains to be seen. While, up to this point, switching to a faster life history has been successful in compensating fitness losses, this strategy requires sufficient habitat quality and is likely risky in times when extreme weather events are becoming more frequent, as predicted by most climate change scenarios.
In times of recent climate change, mechanisms to deal with different environments (e.g. via dispersal to other habitats, or via in-situ responses such as genetic adaptation or phenotypic plasticity) are essential. In regions showing seasonality, organisms are already adapted to regular and, thus, often predictable environmental changes. One well-known strategy to survive periods of food shortage, especially during the winter, is hibernation. Although hibernation is already an adaptation to overcome unfavourable conditions, the optimal timing of hibernation to match for example food abundance peaks is likely to be influenced by changing climatic conditions, as expected during human-induced global change. Thus, the ability to respond to changes in optimal timing of hibernation can be crucial for organisms. All hibernators are positioned at the slow end of the slow-fast life history continuum. Longevity combined with a low annual reproductive output can result in slow recovery from population crashes and is expected to be associated with slow genetic adaptation. Therefore, it is assumed that phenotypic plasticity, a rather rapid and sometimes reversible process, is a crucial mechanism in long-lived organisms to adapt to changing environments. However, how differences in individual hibernation behaviour influence mortality and whether individuals are plastic with respect to their hibernation behaviour are largely unknown.
Recent studies suggest that climatic change can influence hibernation behaviour in various species differently, in a positive or negative way. Female Columbian ground squirrels (Urocitellus columbianus) delayed their emergence from hibernation with later snow melt and lower spring temperatures. Next to the environmental impact, emergence date showed a moderate heritability in female Columbian ground squirrels. Yellow-bellied marmots (Marmota flaviventris) emerged earlier from hibernation with warmer spring temperatures which resulted in a longer growing period for their offspring and, therefore, higher survival rates. In contrast, in alpine marmots (Marmota marmota) lower snow cover due to higher temperatures and, thus, less isolation led to lower juvenile survival. Negative effects, such as reduced juvenile survival, would be of high concern, especially for long-lived species with a low reproductive output.
Bats are exceptionally long-lived compared to other mammals of the same size and often show a low reproductive output with one offspring per year. This is especially true in the temperate zone where bats, furthermore, are characterized by seasonality and depend on hibernation during winter period to survive food and water shortage. Because bats are of high conservation concern it is of prime importance to understand their ability to respond to different climatic conditions and associated mortality costs.
The basis of this study was a five-year data set of 1047 RFID-tagged individuals from two bat species, Natterer’s bats (Myotis nattereri) and Daubenton’s bats (Myotis daubentonii), that were automatically tracked when entering or leaving the joint hibernaculum, “Brunnen Meyer”, located in north-western Germany. The two species are similar sized, share demographical traits and often occupy the same areas. Nevertheless, they differ in their foraging strategy and activity pattern during hibernation period. Natterer’s bats are able to glean insects from surfaces, even at low temperatures. Daubenton’s bats depend on flying arthropods and, thus, warmer temperatures. And indeed there is evidence that Natterer’s bats are able to hunt during hibernation period, while in Daubenton’s bats a lack of feeding during the hibernation period is suggested. Furthermore, Natterer’s bats are characterized by a higher activity at the hibernaculum throughout the hibernation period, while Daubenton’s bats on average arrive earlier, stay inactive through the winter and leave later in spring.
In both species, the aim was to investigate the impact of their individual hibernation behaviour, precisely the timing of departure in late winter and early spring, on mortality, their adjustment of departure timing to the North Atlantic Oscillation Index (NAO), as well as differences within and between the two species from 2011 until 2015.
To later on estimate the potential mortality costs of departure timing, gaining knowledge about the seasonal survival pattern (winter vs. summer) in the two species was a first necessity. In birds, particularly small species were described as winter-regulated populations with a higher mortality during winter. In contrast, in hibernating mammal species, such as bats, a relatively lower or similar winter survival compared to summer survival was shown. In this study, the analysed data demonstrated that the winter 2010/2011 was exceptionally catastrophic in Natterer’s bats and did not impact Daubenton’s bats. When excluding this catastrophic winter in Natterer’ bats, our results revealed a stable winter-summer-survival difference (higher winter and lower summer survival) in adult Natterer’s and Daubenton’s bats, with inter-annual variation in the level of survival which indicates a potential environmental impact on survival. This winter-summer survival pattern is in line with the survival pattern shown for other hibernators. Juveniles always had a lower survival rate than adult bats in both species. Nevertheless, the extent to which the species differ between seasons and age classes was stronger in Daubenton’s bats. They always showed a slightly higher winter survival and a lower summer survival than Natterer’s bats. Together with the catastrophic winter 2010/2011 in Natterer’s bats, this indicates a species-specific sensitivity to the timing of specific weather events which is in line with their foraging strategies and activity pattern during hibernation period.
With respect to emergence behaviour from the hibernaculum, the results of this study suggest considerable differences among individuals within as well as between bat species. In comparison to Daubenton’s bats, Natterer’s bats tuned their emergence more closely to weather conditions, specifically the NAO, a large scale weather index related to winter severity, and showed individual variation in behavioural plasticity. In Daubenton’s bats only the females responded to changing conditions and left earlier in individually-experienced warmer and milder winters, comparable to Natterer’s bats females. A potential reason might be reproductive advantages for the females resulting in a longer growing period for their offspring. The shown higher winter survival in adult bats of both species indicated already higher energy expenditure outside the hibernaculum. Thus, leaving early, being active and staying outside longer by itself bore a risk (exposure risk effect). Under consideration of longer exposure times, early departing individuals had on top of that an increased risk to die. This was not given in each year, but a species- and year-specific pattern was revealed. Natterer’s bats were only significantly affected by early departure in 2011, while the remaining years show no significant additional risk of leaving early. In Daubenton’s bats, the years 2014 and 2015 were associated with a significantly higher mortality of leaving early. This is in line with the hypothesis that Daubenton’s bats might not be able to hunt for insects leaving too early and do so as a best out of a bad job. Nevertheless, the year-specific pattern suggests that early bats might profit from advantageous weather conditions during early spring.
An additional hint for an environmental impact on early bat survival in at least Daubenton’s bats is that the median proportion of night hours above 3 °C within five days after departure was included in the model with the lowest AIC. However, the effect was not strong enough to be selected as the best model and, therefore, further analyses are needed to investigate this first hint.
In conclusion, the reduced winter survival of juveniles compared to adults highlights the importance of considering age class effects in studies that investigate seasonal survival patterns. The stable species-specific winter-summer-survival difference with a higher winter survival compared to summer survival, as well as the one catastrophic winter in Natterer’s bats underline the importance of including seasonal survival patterns in assessing potential fitness costs of changed behaviour. Furthermore, our results suggest that long-lived hibernating bat species have the potential to plastically adjust to changing climatic conditions, but this potential differs between species. Among-individual differences in emergence together with species-specific mortality costs of early emergence suggest the potential for natural selection to shape hibernation phenology. In summary, our findings suggest species-, population- and group-specific differences in the ability to respond to changing environments and, therefore, underline the necessity to further investigate local responses in various organisms to estimate consequences of recent climate change on a wider range.
Recent climate change and its consequences for living organisms constitute one of the greatest problems of our century. Global warming entails an increase in mean temperature and the frequencies of extreme weather events. Those changes in environmental conditions affect both plants and animals. Because of their inability to escape from unsuitable environments, plants have evolved a wide spectrum of molecular programs to protect themselves against changing conditions. Responding on altered environmental conditions will change plants chemical composition and therefore also affect plants interaction with other species (e.g., predator-prey or symbiotic relationships). For instance, changes in the chemical composition of plants may influence the survival of associated herbivores. In other words, these herbivores will be affected indirectly by climate change due to changes in the suitability / quality of their food. The aim of this doctoral thesis was to discover the effects of climate change within the relationship of the butterfly Pieris napi and its host plant (Sinapis alba used here as host plant), including individual conditions (e.g. chemical compositions of plants; morphology, physiology of the butterfly) and behavior of female butterflies and larvae. In the first experiment, the influence of simulated climate change on the chemical composition of the plant Sinapis alba was investigated. The second experiment aimed to examine the influence of changes in plant composition on the butterfly P. napi. Glucosinolates (secondary compound of plants) are known to have an important effect on the preference and performance of herbivores. Therefore, in the third experiment, the impact of glucosinolates on the preference and performance of P. napi was investigated in order to see if these plant compounds had the most important influence on this butterfly. Furthermore, in the fourth experiment, it was explored whether there is a latitudinal gradient within the species´ responses to changes in its host plant. The fifth and last experiment aimed to examine, if there are general principles across species regarding indirect effects of climate change.
Climate change, simulated by different combinations of temperature and water regimes, had an effect on the plant chemistry. The combination of temperature and water availability changed plant composition substantially. Especially the amount of carbon and glucosinolates (here above all sinalbin) in S. alba plants varies between the different treatments and therefore between the different combinations of temperature and water regimes. Regarding glucosinolates, elevated temperatures increased their concentration in leaves, whereas water deficit in combination with higher temperature reversed this pattern. For carbon content, all plants, except those of the control group, showed a decreased amount of total carbon. However, simulated heat waves had no effect on plants, leading to the assumption that the plants were able to recover from heat stress sufficiently during the control phases. Changes in plant composition affected both larvae and females of the butterfly P. napi. Therefore, changed host-plant chemistry alters the plant quality for this herbivore, meaning that plants of different treatments represent different plant qualities defined by their composition. Females of P. napi may be able to differentiate between plant qualities and even show a direct preference. Therefore, glucosinolates seem to act as oviposition stimulants. However, preferring another plant quality with lower amount of glucosinolates suggest that females of this butterfly species were attracted by more than high levels of glucosinolates alone. Larvae fed with different plant qualities performed differently, indicated by smaller wings (lighter bodies) and prolonged development when fed with plants contained higher amount of the glucosinolate sinalbin. It can be assumed that a higher amount of sinalbin decreases the quality of the host plant and therefore lead to these responses. Probably larvae need to shift their resources from growth to detoxification and therewith survival. Furthermore, drought conditions during plant growth seem to reduce the overall negative effects of higher temperatures, lead to an increase of host plant quality. Larvae seem to benefit from feeding on these “double-stressed” plants. Comparison between the results of the preference and performance tests suggests that there might be a mismatch between female preference and larval performance. It seems that the stimulating effect of high concentration of glucosinolates, in this case sinalbin, misdirects females´ decision to less suitable host plants, meaning that the advantage of less competition for larvae come at costs through detoxification. Using Brassica napus plants with genetically fixed glucosinolate levels, it could be demonstrate that there must be other plant components influencing females´ oviposition behavior been seen in the choice experiment with S. alba. The comparison of German and Italian populations to changes in host-plant quality showed fewer differences between countries as expected. However, German and Italian individuals differed in their reaction to altered plant quality, at least in developmental time and larval growth rate. It seems that Italian larvae benefitted from plants grown under higher temperatures, whereas drought-stressed plants affected them negatively. German individuals in contrast seem to benefit only from water stress during plant growth. With regard to the sexes of P. napi, it seems that females respond differently than males to changes in plant quality. Furthermore, the results of the performance test on Bicyclus anynana showed that there might be some general principles for the respond of butterflies to changes of its host plant. B. anynana responded in a similar way to different host plant qualities as P. napi did, meaning that plants grown under higher temperatures and drought conditions seem to be beneficial for the larval performance.
In summary, these findings may have important implications for the indirect effects of climate change on this butterfly in natural environments. First, climate change seems to have an impact on the chemical composition of plants. Second, changes in plants caused by increasing temperature and droughts seem to influence the preference and performance of this butterfly. However, there are differences between populations, which seem to be induced by former adaptation. And third, there might be some general principles for the respond of butterflies to changes in their host plants. This thesis focuses only on possible indirect effects of climate change. However, there are direct effects, which may alter the responses of herbivores to changes in their host plant as well. Therefore, further investigations in this linkage and in other plant-herbivore relationships will be necessary to explore how climate change may alter the relationship between herbivores and their hosts.
Global climate change is omnipresent all over the world and is affecting and challenging organisms in various ways. Species either have to adapt to the changing environmental conditions or move to new habitats in order to avoid extinction.
Possible ways for an organism to react can be dispersal, phenotypic plasticity, genetic adaptation or a combination of these factors. Among the various consequences of climate change, especially changes in temperature affect plenty of species. In ectotherms, the body temperature and associated mechanisms are strongly dependent on environmental conditions.
The aim of this work was to investigate the mechanisms underlying adaptation to thermal variation and heat stress in the widespread butterfly species <i>Pieris napi<i>.
Focusing on indicators of individual condition, including morphology, physiology and life history traits, the purpose was to specify whether the species’ responses to temperature variation have a plastic or genetic basis. In the first experiment, phenotypic variation along a latitudinal and altitudinal cline was investigated. Yellow reflectance of wings was negatively correlated with wing melanisation, providing evidence for a trade-off between a sexually selected trait (yellow color) and thermoregulation (black color). Body size decreased with increasing latitude and led to the assumption that warmer conditions are more beneficial for <i>P. napi<i> than cooler ones. An increased flight performance at higher altitudes but not latitudes may
indicate stronger challenges for flight activity in high-altitude environments.
The second experiment focused on clinal variation and plasticity in morphology, physiology and life history in F1-generation individuals reared in captivity at different temperatures. It could be shown that individuals from cooler environments were less heat-tolerant, had a longer development but were nevertheless smaller, and had more melanised wings. These differences were genetically-based. Furthermore, it could be shown that a higher developmental temperature speeded up development, reduced body size, potential metabolic activity, and wing melanisation but increased heat tolerance, documenting plastic responses.
In a third experiment, we examined physiological responses to heat stress. A transcriptome analysis revealed an upregulation in molecular chaperones under hot conditions, whereas antioxidant responses and oxidative damage remained unaffected. The antioxidant glutathione (GSH) though was reduced under both cold and hot conditions. Interestingly, Swedish individuals were characterized by higher levels of GSH, lower early fecundity, and lower larval growth rates compared with German or Italian populations, suggesting a ‘pace-of-life’ syndrome. Thus, the individuals from warmer regions show the opposite pattern with a lower investment into maintenance but a faster lifestyle.
In summary, we found clinal variation in body size, growth rates and concomitant development time, wing aspect ratio, wing melanisation and heat tolerance. The effects of high developmental temperature very likely reflect adaptive phenotypic plasticity. When speeding up development; heat tolerance is increasing while body size, potential metabolic activity and wing melanisation are decreasing. Overall, body size of <i>P. napi<i> individuals decreased from south to north while the melanisation of the wings increased. Furthermore, we found a connection between increased wing melanisation and decreased yellow reflectance, most likely caused by a trade-off between the two. We could confirm that <i>P. napi<i> individuals from warmer environments were more heat-tolerant and larger than individuals from colder environments. Due to increasing temperatures and heat waves becoming more frequent in the future, being able to cope with such conditions will be advantageous. As warmer conditions had positive effects on individual development, <i>P. napi<i> may benefit from global warming, but its association with moist habitats suggests negative consequences of climate change. We could also reveal pronounced plastic and genetic responses in <i>P. napi<i>, which may indicate high adaptive capacities. Thus, increasing temperature may not be too problematic for the species, as it seems to be rather well equipped to deal with such challenges. However, as climate change entails changes in precipitation / humidity along with temperature changes, such issues need further investigation.
Unstable environments and habitats changing due to climate change force individuals to either respond by genetic adaptation, phenotypic plasticity or by dispersal to suitable environments. Theodoxus fluviatilis (Linneaus, 1758) is a good study organisms when researching phenotypic plasticity and genetic adaptation as it naturally appears in freshwater (FW) as well as brackish water (BW) and thus inhabits a wide range of environmental salinities (0-18‰). It is a euryhaline snail that can be found in shallow waters with stony ground or on Fucus spp. and has formed regional subgroups. The brackish water and the freshwater subgroups are spatially separated and the species cannot be found in areas inbetween, e.g. estuaries.
The species shows great variability in shell patterning and shell size and there is still debate whether the subgroups are distinguishable by these traits or not. The mitochdrial RNA marker cytochrome c subunit I did not show differences between the subgroups indicating that they must be closely related, but salinity tolerance has been observed to be higher in BW snails. This might be caused by the different protein expression patterns and osmolyte accumulation (measured as ninhydrin-positive substances) observed in this species in previous studies. The exact mechanisms regulating protein expression and osmolyte accumulation, however, are not fully understood yet.
Data collected for this thesis shows differences in shell size and suggests a less strict grouping of FW and BW individuals as shell sizes of one FW site are more similar to BW individuals than the other FW ones. A better salinity tolerance towards high salinities and a higher physiological salinity limit of BW snails was confirmed and extended by demonstrating an expanded tolerance range through slow acclimation to challenging salinities in snails from both subgroups. This was achieved by a shift in the slope of their reaction norms that was much more pronounced in BW snails than FW ones. S3 individuals showed a shift similar to that of BW individuals. The data for the salinity tolerance indicates that the underlying mechanism for these tolerances are a combination of phenotypic plasticity and genetic adaptation. Despite an acclimation and shift in the slope of the reaction norms and therefore an increased tolerance towards high salinities (plasticity) FW individuals from two collection sites were not able to cope with salinities as high as BW individuals (local adaptation). The general ability to mobilise free amino acids (FAA) as organic osmolytes was not the reason for this tolerance difference. Individuals from BW and FW sites were capable of accumulating quantities of FAAs equally well. Proline, alanine and urea were the most important components of the accumulated cocktail of organic osmolytes. Even though the total amount of FAAs accumulated under hyperosmotic conditions was the same in both subgroups, there were differences in the metabolic pathways involved in osmolyte accumulation in the foot muscle. The data indicates that the hydrolysis of storage proteins and the synthesis of proline and alanine are the main processes to avoid detrimental body volume shrinkage in T. fluviatilis. While FW individuals seemed to rely on the degradation of proteins and synthesis of alanine, BW individuals depended on newly synthesising proline and alanine and accumulating urea as a side product of transamination. The accumulation of urea is a new finding in aquatic living snails and has not been reported as a mechanism to avoid cell volume shrinkage in these animals.
Differing protein expression patterns were observed under control conditions across all collection sites. 9 spots showed volume changes in BW snails opposite to those of FW snails from collection sites S1 and S2. For 6 of those spots, S3 individuals showed patterns similar to those of BW individuals and for the remaining 3 they showed patterns similar to those of FW animals. The patterns observed when exposing snails to hypo- or hyperosmotic stress were not conclusive in relation to pinpointing individual spots that show the same pattern in all collection sites, but revealed the heterogeneity of protein expression in snails from the different collection sites and in the process of osmoregulation. It also showed the general tendency of protein reduction when snails where under osmotic stress of either kind (hypo- or hyperosmotic), which supports the hypothesis of storage protein degradation.
The investigation of an ANP-receptor showed two variations of the encoding sequence expressed in T. fluviatilis. S3 individuals as well as BW individuals were found to express one type, while FW individuals, with the exception of one sample expressed the other type. This showed that the FW subgroup of T. fluviatilis seems to be more heterogeneous than the BW subgroup, but also raises the question of the dispersal history of this species. The collected data indicates that T. fluviatilis individuals are firstly capable of surviving the acidity of a duck's gizzard and secondly can tolerate acute salinity changes to 16‰ when introduced into a new environment. Hence, if snails from the FW were to be transported to waters with a salinity of up to 16‰ by man, bird, drifting plants or some other means of transport, they would most likely survive and possibly be able to thrive and spread.
Under natural conditions, most parts of northeastern Germany would be covered by forests that would be dominated by beech (Fagus sylvatica) and oak (Quercus robur and Q. petraea). However, today most of the wooded area is covered by artificial monocultures of pine forests. This form of cultivation was recognised to be the cause of instability against calamities of pests as well as severe storms therefore in the early eighties of the last century this knowledge was used to start the conversion of the forests towards more nature-like stands. The ecological effects of the forest conversion on the soil, the fauna and the flora have been investigated in a nation-wide project supported by the Federal Ministry of Education and Research (BMBF) in the project “Future-oriented forest management”. The present work has been accomplished within the scope of this project and is concerned about the effects that different aspects of forest conversion have on oribatid mites. The present work shall serve to answer a number of questions about the distribution of oribatid mites and their reaction to environmental changes. The investigation was carried out on 12 plots in two sampling areas. 7 plots were chosen in the Müritz NP and 5 in Eberswalde. In both areas plots were chosen that resemble the different stages of forest conversion: one medium aged pine plot in each area, two medium aged mixed plots with pines and beeches in the Müritz NP and one mixed plot in Eberswalde as well as one beech plot in each area. Furthermore, in the Müritz NP the chance arose to investigate the effects of different age stages of the stands on the oribatid mites. Therefore, an additional young pine plot and two old mixed plots have been sampled. In Eberswalde, on the other hand, another emphasis was laid on the effects of a different nutrient content in the soil. Here, an additional pine plot and mixed plot, respectively, of a higher trophotopic level was sampled. In Eberswalde, an additional sampling was done in three plots (a beech plot, a mixed plot and a pine plot) to investigate the horizontal distribution of the oribatid mites in these habitats. The data were used along with others to ecologically characterise the different species. The sampling took place from 2000 to 2002. Within the scope of the doctoral thesis, 392 samples were analysed. 122 samples from one year from the Müritz NP and 270 samples from three years from Eberswalde were analysed. Altogether 155,450 oribatid mites from 82 taxa were found in these samples. The ecological characterisation of the species revealed that the various species react quite differently to the investigated factors. Most species occur with different abundances in different forest types, but their abundance often varies also in comparable stands of both sampling areas. This indicates that they react to climatic effects as well as to biotic and abiotic factors. The forest conversion from pine forests to beech forests causes the abundance of oribatid mites to decrease, probably due to the change of the humus form from mor or mor-like moder in pine forests to mull in beech forests, that is accompanied by a decrease of the abundance of fungi, the main food source for most oribatid mites. Furthermore, the species composition changed. Species like Camisia spinifer, Adoristes ovatus or Acrogalumna longipluma that are typical for pine forests disappeared, while other species like Achipteria coleoptrata or Chamobates voigtsi immigrated in mixed stands after the introduction of beeches. The age of the stands proved to be another important factor. The overall abundance of oribatid mites was higher in the older stands than in the younger stands, while the percentage of juvenile oribatids decreased towards the older stands. Furthermore, the dominance structure became more uneven and shifted toward a higher percentage of fungivorous oppiid and suctobelbid mites. Especially on the old mixed plots, Oppiella nova reaches a dominance value of about 60 %. The nutrient content of the soil seems to be a relatively unimportant factor on the community level as no significant differences with regard to overall abundance and the dominance structure could be recorded. However, the Canonical Correspondence Analysis showed that the nutrient content of the soil does influence the distribution of species, at least with regard to their individual abundance. In summary, it can be said that the distribution of the oribatid species is influenced by many factors, and the stocking is only one of these factors. Nevertheless, a group of four species could be established, that can be used as indicators for the success of the forest conversion towards more nature-like deciduous forests: Achipteria coleoptrata, Autogneta longilamellata, Chamobates subglobulus and C. voigtsi.
In holometabolic insects, senescence starts at sexual maturation and condition diminishes with age. Young virgin males should gain the highest mating success. Although sperm quality and quantity typically decreases with age, older males have been shown to have a higher mating success in a variety of insect taxa. Life-history theory predicts an increased aggressive and persistent courtship behaviour, due to a decrease of the residual reproductive value and thus, the opportunity for future reproduction. In the butterfly Bicyclus anynana, older males gain up to a 4x higher mating success despite a poorer condition, compared to younger males. As older males were found to court more often and for longer time bouts,suggesting a higher intrinsic motivation, there is evidence for the residual reproductive value hypothesis. On the other hand, an age-specific variation of male sex pheromones may provide females with information, being helpful to chose specific males. The latter could be males which have proven a high fitness in terms of survival, thus essentially reflecting a “good genes" hypothesis. This doctoral thesis investigated the reasons underlying old male mating advantage in B. anynana. The first study tested whether old male mating advantage prevails, even if females were unable to distinguish between older and younger males. The results were backed up by examining female rejection rates, male courtship frequency, courtship duration, time to copulation and mating duration. Older males had a significantly higher mating success compared to younger ones regardless of differences in pheromone blends or the females ability to smell. Older males courted more often and longer. The results support the residual reproductive value hypothesis. Study 2 investigated post-copulatory sexual selection in B. anynana. Females were double-mated, allowing for sperm competition and cryptic mate choice. Older males had a higher paternity success than younger ones, when the former were the last mating partner. The paternity success was balanced across age classes when older males were the first mating partner. Older males transferred larger spermatophores with higher numbers of eupyrene sperm than younger males. B. anynana does not exhibit cryptic mate choice and last-male precedence is the most probable fertilization pattern. The higher proportion of offspring sired by older males is due to significantly increased numbers of fertile sperm compared to younger males. The latter are clearly disadvantaged, as the higher sperm numbers provided by older males counteract the benefits of last-male precedence. Study 3 explored determinants of mating success in B. anynana. Successful males had longer wings, a heavier thorax, a lighter abdomen, a higher fat content, and higher phenoloxidase expression levels. Mating success seemed to be directly linked to proxies of flight performance. Successful males showed a better flight performance, in turn increasing mating success. As energy reserves are of crucial importance for flight manoeuvres, they may play a significant role in male mating success. Study 4 tested the effects of male age and mating number on spermatophore mass, sperm number, male oxidative status, and reproductive success by manipulating age and mating frequency in male B. anynana. Spermatophore mass and sperm numbers increased significantly with age, while antioxidant defences and oxidative damage declined. Female fecundity and egg-hatching success was highest when being mated with young virgin males. Antioxidant defences decreased with age, being a possible reason for the negatively affected reproductive success in females. Ejaculate quality diminished with age and mating number, despite larger spermatophores and higher sperm numbers being found in older males. Therefore, spermatophore size and sperm numbers can not be considered as reliable proxies of male condition. The final study proofed whether male sex pheromone comprise honest signals and which traits might be associated with increased pheromone titres. Pheromone titres were analysed among successful and unsuccessful males when being mated with either a control or a scent-blocked female. Both groups did not differ in pheromone levels. Successful males had significantly higher numbers of eupyrene sperm. No correlation between male pheromones and any investigated trait was found, suggesting that pheromone titres do not provide reliable information on male quality. Nothing indicates that male sex pheromones in B. anynana play a decisive role in female mate choice. Successful males generally have a better body condition, resulting in a more vigorous courtship behaviour and higher sperm numbers. Hence, variation in body condition rather than pheromone titres is more likely to determine male mating success. Mating decisions are primarily driven by male behaviour. The results suggest that old male mating advantage arises from sexual conflict.
Decision making in everyday purchase situations requires mental processing of factors that are related to the items on display. These influencing factors – called persuasive information – can take various forms, like the price level, the design of the package or the display of certain product attributes. Despite the existence of persuasive information trying to influence our buying behavior, almost nothing is known about the underlying neural mechanisms responsible for processing this information. In this thesis functional magnetic resonance imaging was used to investigate neural activity correlated with product related persuasive information. As persuasive information organic, light and regular labeled food was chosen. The 1st experiment investigated the neural correlates of visually inspected organic and regular labeled food and the influence on willingness to pay (WTP) for the displayed items. It was hypothesized that organic compared to regular labeled food will be perceived as more rewarding which should be visible by an increased activity in the ventral striatum as a central area for reward processing and by a heightened WTP. As organic label information the national German eco emblem 'Bio-Siegel' was chosen (for stimuli details see 2.1). As there is no emblem indicating regular food, an artificially created logo was used for indicating a conventional product. 40 well- known food products (e.g. milk, bread, eggs etc.) were presented to the subjects. These products were marked with the organic emblem and the same 40 products with the regular label. We found that visual inspection of organic labeled food indeed led to an increase in neural activity in the ventral striatum and to a heightened WTP, suggesting a higher subjective value for these products. The 2nd experiment investigated the neural correlates of actually administered food stimuli labeled organic, light or regular and the influence on expected and experienced taste. For organic compared to regular labeled food we hypothesized an increase in expected and experienced taste pleasantness. Furthermore, light compared to regular labeled food should lead to a decrease in expected and perceived pleasantness and intensity ratings. During the active tasting process this should be accompanied by an increase in reward-related (e.g. organic vs. regular; regular vs. light) areas like the ventral striatum medial orbitofrontal cortex or aversion-related (e.g. regular vs. organic; light vs. regular) areas as the lateral orbitofrontal cortex (lOFC) and operculum/insula. As organic label information the national German eco emblem 'Bio-Siegel' was chosen. Light label information was issued in form of the internationally used 'Bewusst-Wählen®' ('Healthy Choice') label (for stimuli details see 2.2). However, inside the scanner the written forms 'Bio', 'Light' or 'Normal' (indicating regular food) were chosen. Subjects were randomly assigned in two groups and were either confronted with the organic or the regular label (organic group) or with the light or the regular label (light group) but otherwise identical milk drink. The results show that organic compared to regular labeling of identical food stimuli indeed led to an increase in expected and experienced taste pleasantness for organic labeled food. Light compared to regular labeling of identical food stimuli led to a decrease in expected and experienced taste pleasantness and intensity for light labeled food. Moreover, taste-related activity was found in aversion related areas like the operculum insula and the lOFC for food labeled regular compared to organic and in reward-related areas like the ventral striatum for food labeled regular compared to light. The results show that persuasive food-related information influences human cognition on the behavioral and neural level; the effects were shown during visual and gustatory evaluation of the stimuli. Taken together the results demonstrate that the same stimulus can vary dramatically in personal valuation depending on the applied information.
The goal of this thesis was to study the systematic relationships within the superfamily Sylvioidea (Aves: Passeriformes) in general and within the closely related families Acrocephalidae and Locustellidae in particular, by means of DNA sequences. Sylvioidea itself and families therein were the focus of many studies based as well on morphological characters as on DNA. Due to their morphological similarity and their presumably rapid radiation most studies failed to solve relationships between sylvioidean families and also demarcations of single families and relations within are still in progress. In this study, an enlargement of previous datasets, both taxa and number of DNA sequences, and more sophisticated analysis methods were used to improve the resolution in Sylvioidea, Acrocephalidae and Locustellidae. In addition, the applicability of barcoding in Acrocephalidae was tested. The monophyly of Sylvioidea could be supported and the families Paridae and Remizidae, which were sometimes still included, clustered among the outgroup taxa. Four families, Nicatoridae, Panuridae, Alaudidae, and Macrosphenidae constitute basal splits within Sylvioidea. The division of the former sylviid/timaliid clade in five families, Sylviidae, Leiothrichidae, Pellorneidae, Timaliidae, and Zosteropidae was supported. Scotocerca, Erythrocercus, and Hylia, previously supposed to be members of Cettiidae, were shown not to belong to this family. As the three genera are also morphologically and ecologically different, they were here proposed to be elevated to family rank, Scotocercidae, Erythrocercidae and Hyliidae, respectively. The family Acrocephalidae consisted of the four genera, Nesillas, Acrocephalus, Hippolais, and Chloropeta. In the analysis for this thesis, the latter three appeared to be non-monophyletic. One Acrocephalus species, A. aedon was sister to a clade containing four species of Hippolais as well as two out of three Chloropeta species. They were all merged in the genus Iduna, based on the DNA evidence and shared morphological and ecological characters. Iduna had priority over Hippolais or Chloropeta according to the International Code of Zoological Nomenclature. The one remaining Chloropeta species (C. gracilirostris) had to be renamed to Calamonastides as Chloropeta was no longer available for this taxon. Seven genera were included in the re-analysis of the family Locustellidae: Locustella, Bradypterus, Megalurus, Dromaeocercus, Schoenicola, Cincloramphus, and Eremiornis. Apart from the monotypic genera Dromaeocercus and Eremiornis and Schoenicola, of which only one species was included, the remaining genera were found to be non-monophyletic. One clade contained all Locustella species, Megalurus pryeri and all Asian/Oriental Bradypterus species. All species in this clade were synonymized with Locustella, as the type species of Locustella was included, whereas the type species of Bradypterus fell in a different clade. Therefore, the remaining African Bradypterus species retained their genus name, and Dromaeocercus was renamed to Bradypterus as it clustered within Bradypterus. Cincloramphus, intermingling with the remaining Megalurus species, was synonymized with the latter. Barcoding, growing in popularity for delimiting species, was tested in its applicability for Acrocephalidae. Fourteen taxa currently recognized as full species would fall under the 2% threshold of sequence divergence proposed for delimiting species using the mitochondrial cytochrome b gene. It was also shown that it is important to clarify which part of a DNA sequence is used, as different parts can give different results regarding the 2% threshold. In addition, the choice of “complete deletion” or “pairwise deletion” in calculating genetic distances is important, if incomplete are sequences used.
Bats are special: although they have a small body size, bats are extremely long-lived and have a low annual reproductive output, which puts them at the ‘slow’ end of the slow-fast continuum of mammalian life-histories. Species typically respond to climate change by genetic adaptation, range shifts or phenotypic plasticity. However, limited dispersal behavior in many bat species and long generation times make it very likely, that adaptive responses in bats are rather driven by phenotypic plasticity than by genetic adaptation or range shifts. Changing weather patterns, a higher frequency of extreme weather events and overall rising temperatures, caused by climate change, will impact phenology, energy supply and energy expenditure. In species where adult survival largely shapes population dynamics, it is thus of crucial importance to understand how climate change affects individual fitness and fitness relevant traits by altering behavior and development.
In my study, I investigated how weather impacts behavior, fitness and fitness relevant traits in free ranging Natterer’s bats from two geographical regions (south vs. north) in Germany. In the Nature Park Nossentiner/Schwinzer Heide (northern region, NSH), long-term data for investigations on population dynamics are partially collected by hibernation counts. Although counting hibernating bats is a regularly applied method, it is still unclear to which degree human visits in the hibernaculum trigger energy consuming arousals and thus increase energy expenditure. Thus, I first investigated if hibernation counts potentially threaten winter survival by assessing the number of energy consuming arousals of hibernating Natterer’s bats (Myotis nattereri) and two other bat species (Pipistrellus spp., Plecotus auritus) using thermal imaging. Additionally, I used light barriers in the hibernacula to investigate the relative impact of winter temperatures and human visits on flight activity of hibernating bats. Secondly, I investigated effects on survival and reproduction during summer by analyzing capture-mark-recapture data from summer roosts. Data from summer roosts have been collected since 2011 in Würzburg (WB, south) and 1990 in the Nature Park (NSH, north). Based on these data, I analyzed the effect of intrinsic (e.g. age) and extrinsic(e.g. different weather parameters) factors on individual survival probability and reproductive success. I further focused on the question if individual body size is a fitness relevant trait in Natterer’s bats and how body size of young bats is affected by summer temperatures.
During hibernation, ambient temperatures were the most important driver for bat activity and were positively correlated with the number of flight passes in the light barrier, suggesting that bats can exploit foraging opportunities more frequently during warm weather bouts. Monitoring caused only a small number of arousals and only a slight increase in activity, which was less severe on warmer days, when activity was generally higher. Thus, I propose that benefits of hibernation counts outweigh the costs of human presence in the hibernaculum and unlikely threaten winter survival in hibernating bats.
In spring, increased precipitation during a short time window strongly reduced the probability of successful reproduction in first-year females (females that returned from first hibernation, FY). In terms of timing, this sensitive period comprises the implantation or early pregnancy, a time before substantial investment into embryo development. Moreover, I identified a positive correlation between a large body size and reproductive success in FY females. Given the evidence that suitable weather conditions during early life support juvenile growth and thus a large body size, my findings suggest that reproduction may be condition dependent in young females. Reproductive success of older females was not affected by either weather or individual parameters. This suggests that older and experienced females can better deal with adverse conditions.
To examine if beneficial weather conditions are linked to a large body size, I investigated the effect of ambient temperatures during the growing season on body size. I found that higher ambient temperatures during summer led to larger individuals, however, only in the northern population. In the on average colder North, warmer summers may benefit juvenile growth by reducing thermoregulatory costs and increasing prey abundance, whereas in the general warmer South, this effect might not be visible or relevant. When I analyzed the link between body size and survival, I revealed that larger adult females have higher survival rates. Given the fact, that a large body size is a response to beneficial early life conditions, this demonstrates the impact of early life conditions on long lasting fitness effects.
The results of my research lead to the assumption that warmer ambient temperatures have positive effects on Natterer’s bats, both during winter and summer. However, increased activity in response to rising winter temperatures, as expected under climate change scenarios, could be a serious thread for hibernating bats, if food availability does not increase in the same amount as bat activity. During summer, warmer temperatures may have positive effects on juvenile development in northern regions, but this effect could be negative in more southern regions by exceeding heat tolerance and resulting in water stress. This research highlights, that investigating periods of weather sensitivity on a finer time scale and also in a spatial context is of crucial importance to gain a better understanding for mechanisms leading to the impacts of weather on fitness.
The impact of inbreeding under different environmental conditions and of artificial selection on cold tolerance was investigated in laboratory populations of the tropical butterfly Bicyclus anynana. The investigation focused on (1) the effects of inbreeding on several fitness-related traits and whether inbred individuals are more susceptible to stress, (2) interactions between inbreeding, genetic adaptation to cold stress and environmental conditions, (3) the effects of artificial selection and inbreeding in the adult stage in other developmental stages, and (4) the effect of inbreeding depression on the heat shock response. Environmental conditions are not constant over time; consequently organisms have to deal with environmental changes. Besides naturally fluctuating conditions, human-induced climate change may increase temperature changes as well as the severity of heat or cold waves. Temperature-stress resistance describes an organism’s ability to cope with stressful temperatures. Enhanced resistance to temperature stress can be reached by phenotypic plasticity or genetic adaptation. Plastic organisms are able to react fast to changing environmental conditions, whereas genetic adaptation is more important for long-term adaptation. Natural habitats may also be affected by human impact, causing habitat loss or fragmentation and changes in population structure. A decrease in the population size may result in inbreeding and inbreeding depression (ID). Consequences of inbreeding are well documented, and inbred individuals are predicted to be more sensitive to environmental stress than outbred individuals. The long term persistence of species and populations depends on their ability to adapt to novel conditions which in turn depends on genetic diversity. Therefore, studies of temperature resistance and its evolution in relation to inbreeding are very important. First a higher susceptibility of inbred individuals to environmental stress was determined in different populations of B. anynana. Inbreeding depression was revealed for several fitness-related traits, but not for immunity traits or heat tolerance. Temperature affected most traits, revealing the importance of temperature on ectotherms; just two hours of thermal stress affected important reproductive, life-history and immunity traits already. Importantly though, no evidence were found that inbred individuals are more susceptible to stressful temperatures than outbred individuals. Genetic adaptation and phenotypic plasticity can interact with one another, resulting in genotype-environmental interactions (G x E). The hypotheses tested here were that some genotypes are more plastic than others and that lines with increased cold stress resistance are less plastic with regard to cold resistance than control lines. To induce plastic responses the exposed lines differed in cold tolerance and inbreeding to different temperatures as well as different feeding regimes and measured fitness-related traits. Several interactions were detected in which a selection regime was involved, but these interactions did not show a clear overall pattern. In summary though, findings were that marginal impacts of directional selection and inbreeding on plastic responses and suggest that, at least for my study organism, the genetic architecture of fitness-related traits is not connected with the architecture of plastic responses. The next investigation concerned with the manifestation of genetic adaptation to produce one specific phenotype across development stages and possible trade-offs. The assumption tested was that there is a genetic link between different developmental stages to produce one definite phenotype by imposing selection in the adult stage only. Lines selected for increased cold resistance in the adult stage were used and increased cold resistance throughout all developmental stages was expected. However, higher cold resistance was found only in the adult stage and not in developmental stages. This could be either the result of a resource allocation trade-off between different stages or that there is no cold resistance phenotype. Thus, if selection takes place in the adult stage it does not affect the others. In the last experiment investigation was directed to determine whether there are negative inbreeding effects on the heat shock protein (HSP) response. Under stressful conditions, organisms produce the HSPs and they act as chaperons required for refolding and repairing of stress degraded proteins. Testing was oriented to find if inbreeding as a genetic stressor´ provokes a higher HSP expression and if there is evidence for higher temperature stress susceptibility on inbred individuals. Findings indeed showed a stronger HSP up-regulation in control compared to inbred lines with a negative inbreeding impact occurrence, which may causally underlie inbreeding depression.
Amid the current global biodiversity crisis, being able to accurately monitor the changing state of biodiversity is essential for successful conservation actions and policy. Despite the pressing need for reliable and cost-effective monitoring methods, collecting such data remains extremely difficult for elusive species, such as temperate zone bats. Although bats are important indicators of environmental changes, monitoring bat populations is challenging because they are nocturnal, volant, small, and highly sensitive to human activities and disturbance. Thus far, population trends of temperate zone bats have been mainly based on visual surveys, including winter hibernation counts at underground sites. However, as bats may not always be roosting in visible locations within the hibernacula, it is currently unknown how these estimates relate to actual population sizes.
Infrared light barriers combined with camera traps are a novel method to monitor bats at underground sites. When installed at the entrance of hibernacula, infrared light barriers have the potential to estimate site-level population sizes more accurately than visual surveys, by counting all bats flying in and out of the site. Moreover, camera traps, consisting of a digital camera and white flash, can be used for species-level identification. However, for this new method to be applicable as a large-scale bat monitoring technique, it is important to characterize it with regard to three main criteria: is the method minimally invasive, is it accurate, and is it scalable in terms of spatial and temporal resolution? Therefore, the purpose of this thesis was to investigate the invasiveness and accuracy of this novel bat monitoring method, and to develop standardized and automated data analysis pipelines, both for the light barrier and camera trap data, to support the deployment of this method at scale.
In Publication I, we used light barrier data, infrared video recordings and acoustic data from an experimental field study to investigate whether the white flash of the camera trap has any measurable short- or long-term effect on bat activity and behavior. The flash of the camera trap was turned on and off every week at each site, which allowed us to compare the activity and behavior of bats between flash-on and flash-off nights. We found that despite the high sensitivity of bats to disturbance, they did not change their nightly activity patterns, flight direction, echolocation behavior, or long-term site use in response to the white flash of the camera trap. Based on these results, we concluded that camera traps using a white flash are a minimally invasive method for monitoring bat populations at hibernacula, providing high quality images that allows species-level identification.
In Publication II, we used infrared video surveillance to quantify the accuracy of infrared light barriers, and we described a standardized methodology to estimate population sizes and trends of hibernating bat assemblages using light barrier data. We showed that light barrier accuracy varies based on the model and location of the installation relative to the entrance, with the best combination achieving nearly perfect accuracy over the spring emergence phase. When compared to light barrier-based estimates, we found that visual counts markedly underestimated population sizes, recovering less than 10% of the bats at the most complex hibernacula. Moreover, light barrier-based population trends showed regional patterns of growth and decline that were not detectable using the visual count data. Overall, we established that the light barrier data can be used to estimate the population size and trends of hibernating bat assemblages with unprecedented accuracy and in a standardized way.
In Publication III, we described a deep learning-based tool, BatNet, that can accurately and efficiently identify bat species from camera trap images. The baseline model was trained to identify 13 European bat species or species complexes using camera trap images collected at 32 hibernation sites (i.e., trained sites). We showed that the baseline model performance was very high across all 13 bat species on trained sites, as well as on untrained sites when the camera angle and distance from the entrance were comparable to the training images. At untrained sites with more atypical camera placements, we demonstrated the ability to retrain the baseline model and achieve an accuracy comparable to the trained sites. Additionally, we showed that the model can learn to identify a new species, while maintaining high classification accuracy for all original species. Finally, we established that BatNet can be used to accurately describe ecological metrics from camera trap images (i.e., species diversity, relative abundance, and species-specific phenology) that are relevant for bat conservation.
We conclude that infrared light barriers and camera traps offer a minimally invasive and accurate method to monitor site-level bat population trends and species-specific phenological estimates at underground sites. Such remote data collection approaches are particularly relevant for monitoring large, complex hibernation sites, where traditional visual surveys are not feasible or account only for a small fraction of the actual population. Combining this automated monitoring method with a deep learning-based species identification tool, BatNet, allows us quickly and accurately analyze millions of camera trap images resulting from large-scale, long-term camera trap studies. As a result, we can gain unprecedented insights into the behavior and population dynamics of these enigmatic species, drastically improving our ability to support data-driven bat conservation.
Solifuges (Solifugae, Arachnida) are an important element of the fauna especially in arid and desert environments. Unfortunately, this animal group has only been extremely poorly studied not only in terms of morphology, but also ecology, physiology, systematics and phylogeny. The present study aimed to provide a detailed overview of their anatomy and ultrastructure. Representatives of these peculiar animals were investigated by means of light and electron microscopy in order to gain new insights in their functional morphology and also to evaluate potential characters for their systematic and future phylogenetic studies. The histology and ultrastructure of the following organ systems have been investigated: tarsal structures, sensory structures and nervous system, coxal glands, alimentary system, respiratory system, circulatory system and reproductive systems. Additionally, a camel spider in Baltic amber was described, representing the second known specimen of fossil Solifugae in Baltic amber. Further on, the entire mitochondrial genome of Nothopuga sp. was sequenced for comparison with other solifuges and chelicerates in order to reveal changes in their gene order.
Under the influence of human activities, increased climate variability induces changes in
multiple marine environments. Especially vulnerable are the coastal ecosystems where organisms
must cope with constant extreme changes of environmental drivers, such as temperature, salinity, pH,
and oxygen content. In coastal areas, brachyuran crabs are important animals that have a high impact
on ecosystem functioning and serve as a link in food webs and pelagic-benthic coupling. Larval stages
of crabs are crucial for population persistence and dispersal. They are generally more vulnerable to
changes of environmental drivers and failure to adapt to new conditions may result in population
collapse. To analyse the effects of multiple environmental drivers on larval performance and to
elucidate interspecific and intraspecific difference, this project examined larval performance in the
European shore crab Carcinus maenas. In this study, larvae of C. maenas from three native
populations (Cádiz: Cádiz Bay, Helgoland: North Sea, Kerteminde: Baltic Sea) were reared in a
factorial design consisting of different temperature (15-24 °C) and salinity treatments (20, 25, 32.5
PSU). Results demonstrated how descriptors of larval performance (growth, physiological, and
developmental rates, and survival) were affected by combined environmental drivers. Larval
responses to temperature and salinity showed contrasting patterns and differed among native
populations originating from distant or contrasting habitats, as well as within the populations. The
highest overall performance was recorded in the Cádiz population, while the Kerteminde population
had the lowest performance in most of tested traits. The interactive effects of multiple drivers differed
among the populations. In the Cádiz and Helgoland populations, higher temperatures mitigated the
effect of lower salinity while the Kerteminde population showed a maladaptive response when
exposed to lower salinity. Differences in performance showed better locally adapted populations (e.g.
Cádiz) that could acclimate faster, have better adaptive mechanisms or stronger dispersive abilities.
Because of their wider tolerance to increased temperature and decreased salinity, interactive effects
in particular populations may favour some populations in a changing climate, especially in coastal
habitats. Variation in larval performance showed complex interactions in larval performance and
highlighted the necessity to quantify inter-population responses to climate-driven environmental
change where responses of species should not be generalised. This study emphasizes the need for
inclusion of multiple traits, drivers, and populations in experimental studies to properly characterize
performance of marine coastal animals.