Der Streifenkiwi (Apteryx mantelli) kommt im Freiland nur auf der Nordinsel Neuseelands vor. Aufgrund des gefährdeten Bestands ist eine sich selbst erhaltene Zoopopulation wichtig. Kenntnisse des Verhaltens helfen, die Ansprüche der Tiere zu verstehen. Zudem können sie darüber Auskunft geben, inwiefern das Wohlbefinden eines Tieres gegeben ist. Durch die Untersuchung der Brutaktivität sollte ein Überblick über den allgemeinen Verlauf der Brut gegeben und Aktivitätsmuster für den Berliner Hahn erarbeitet werden, um den Verlauf zukünftiger Bruten einschätzen und eventuell positiv beeinflussen zu können. Dazu kamen die Untersuchung der täglichen Aktivität einer Henne sowie Beobachtungen des Verhaltens der Tiere. Diese dienten der Bestandsaufnahme der gezeigten Verhaltensweisen und sollten zusammen mit der Aktivität die Grundlage zur Einschätzung bilden, ob die Ansprüche der Kiwis im Zoo Berlin erfüllt werden, und Hinweise zur Verbesserung der Haltung geben. Die Brutaktivität des Hahnes konnte über drei Brutperioden hinweg detailliert dargestellt werden und zeigte, dass nicht nur innerhalb der Art sondern bei einem einzigen Tier unter ähnlichen Bedingungen die Variabilität so groß sein kann, dass sie für Vorhersagen über den Erfolg einer Brut nicht geeignet ist. Im Zusammenhang mit der Aktivität der Henne ließen sich keine Auffälligkeiten erkennen, die auf eine allgemeine Störung der Tiere schließen lassen oder für eine Beeinträchtigung der Brut verantwortlich gemacht werden könnten. Soweit aus den Beobachtungen im Freiland geschlossen werden kann, scheinen die Kiwis im Zoo ein weitgehend natürliches Verhalten zu zeigen. Die Haltungsbedingungen scheinen den Ansprüchen der Tiere zu entsprechen. Es ließen sich nur bedingt Strategien entwickeln, um die Bedingungen für die Brut und damit für die Nachzucht zu verbessern, da sich die Aktivität des Hahnes während der Brut von Jahr zu Jahr als unerwartet variabel erwies. Für ein weiteres Verständnis des Brutverhaltens und eine mögliche Verbesserung der Bedingungen wäre eine Untersuchung zum Einfluss verschiedener Umweltfaktoren auf die Brutaktivität des Hahnes wünschenswert. / The North Island brown kiwi (Apteryx mantelli) is an endemic inhabitant of New Zealand's North Island. A self-preserving population at the zoos is important because of endangerment in the wild. Knowledge about their behaviour can help to understand the animal's requirements. Additionally it can provide information on the extent to which their well-being is given in captivity. Observing the breeding activity should lead to an overview of the brood's characteristics and the male's activity pattern in order to assess future broods and possibly influence them in a positive way. Additional observations about the female's daily activity and the male's and female's behaviour were conducted. This led to an inventory of their behavioural patterns. Together with the activity data, this provided a basis for assessing the well-being of the kiwis at Berlin Zoo and to optimize the keeping conditions. The male's breeding activity was described in detail for three breeding periods. It was shown that the variability was very high under similar conditions, so predictions about the breeding success were feasible. In conjunction with the female's activity there were no indications implying an overall disturbance negatively influencing the animals or their breeding. As far as it is known from observations in the wild, kiwis at the zoo seem to widely display natural behavioural patterns. The keeping conditions appear to meet the animal's demands. There were only few strategies that could be developed for further optimization of the breeding conditions because of the high variability of the male's activity. For a deeper understanding of the breeding behaviour and possible improvement of the breeding and keeping conditions, additional research about the activity's dependency on environmental factors would be desirable.

Die nichtproteinogene Aminosäure GABA (γ-Aminobuttersäure) gilt als der wichtigste inhibitorische Neurotransmitter im Zentralnervensystem von Vertebraten sowie Invertebraten und vermittelt ihre Wirkung u. a. über die metabotropen GABAB-Rezeptoren. Bisher sind diese Rezeptoren bei Insekten nur rudimentär untersucht. Für die Amerikanische Großschabe als etablierter Modellorganismus konnte pharmakologisch eine modulatorische Rolle der GABAB-Rezeptoren bei der Bildung von Primärspeichel nachgewiesen werden. Ziel dieser Arbeit war eine umfassende Charakterisierung der GABAB-Rezeptor-Subtypen 1 und 2 von Periplaneta americana. Unter Verwendung verschiedenster Klonierungsstrategien sowie der Kooperationsmöglichkeit mit der Arbeitsgruppe von Prof. Dr. T. Miura (Hokkaido, Japan) in Hinsicht auf eine dort etablierte P. americana EST-Datenbank gelang die Klonierung von zwei Rezeptor-cDNAs. Die Analyse der abgeleiteten Aminosäuresequenzen auf GB-spezifische Domänen und konservierte Aminosäure-Reste, sowie der Vergleich zu bekannten GB Sequenzen anderer Arten legen nahe, dass es sich bei den isolierten Sequenzen um die GABAB-Rezeptor-Subtypen 1 und 2 (PeaGB1 und PeaGB2) handelt. Für die funktionelle und pharmakologische Charakterisierung des Heteromers aus PeaGB1 und PeaGB2 wurden Expressionskonstrukte für die Transfektion in HEK-flpTM-Zellen hergestellt. Das Heteromer aus PeaGB1 und PeaGB2 hemmt bei steigenden GABA-Konzentrationen die cAMP-Produktion. Die Substanzen SKF97541 und 3-APPA konnten als Agonisten identifiziert werden. CGP55845 und CGP54626 wirken als vollwertige Antagonisten. Das in vitro ermittelte pharmakologische Profil im Vergleich zur Pharmakologie an der isolierten Drüse bestätigt, dass die GABA-Wirkung in der Speicheldrüse tatsächlich von GBs vermittelt wird. Für die immunhistochemische Charakterisierung konnte ein spezifischer polyklonaler Antikörper gegen die extrazelluläre Schleife 2 des PeaGB1 generiert werden. Ein weiterer Antikörper, welcher gegen den PeaGB2 gerichtet ist, erwies sich hingegen nicht als ausreichend spezifisch. Western-Blot-Analysen bestätigen das Vorkommen beider Subtypen im Zentralnervensystem von P. americana. Zudem wird der PeaGB1 in der Speicheldrüse und in den Geschlechtsdrüsen der Schabenmännchen exprimiert. Immunhistochemische Analysen zeigen eine PeaGB1-ähnliche Markierung in den GABAergen Fasern der Speicheldrüse auf. Demnach fungiert der PeaGB1 hier als Autorezeptor. Weiterhin konnte eine PeaGB1-ähnliche Markierung in nahezu allen Gehirnneuropilen festgestellt werden. Auch die akzessorischen Drüsen der Männchen, Pilzdrüse und Phallusdrüse, sind PeaGB1-immunreaktiv. / The non-proteinogenic amino acid GABA (γ-aminobutyric acid) is the major inhibitory neurotransmitter in the central nervous system of vertebrates and invertebrates, and GABA mediates its action among others via metabotropic GABAB receptors (GBs). So far, these receptors are only rudimentary characterized in insects. In the American Cockroach, which is an established model organism, pharmacological studies have pointed out a modulatory role of GBs in the production of primary saliva in the salivary gland. Therefore, the aim of this study is the profound characterization of the GABAB receptor subtypes 1 and 2 of Periplaneta americana. Diverse cloning strategies and the access to a Periplaneta EST-database enabled the cloning of two receptor-cDNAs. The analysis of the deduced amino acid sequences for GB-specific domains and conserved amino acid-residues, and the comparison to known GB-sequences of other species revealed that the sequences correspond to GABAB receptor subtypes 1 and 2 (PeaGB1 and PeaGB2). Next, we functionally and pharmacologically characterized the receptor-heteromer of PeaGB1 and PeaGB2. Therefore, we established expression constructs for the transfection of HEK-flpTM-cells. The PeaGB1/PeaGB2 heteromer inhibits dose-dependently the production of cAMP. The substances SKF97541 and 3-APPA imitate the GABA effect. In contrast, CGP54626 and CGP55845 are considered to be proper antagonists. The comparison of the in vitro with the known pharmacology of isolated glands reveals that GBs indeed mediate the effect of GABA in the salivary gland. For immunohistochemical localization, a specific polyclonal antibody was raised against the extracellular loop 2 of PeaGB1. A second antibody, which was raised against the analogous region of the PeaGB2, must be considered to be non-specific. Western blot analyses demonstrate the localization of both subtypes in the central nervous system of P. americana. Additionally, PeaGB1 is expressed in the salivary gland and in male accessory glands. Immunohistochemical analyses reveal the expression of PeaGB1 in GABAergic nerve fibers of the salivary gland. As a consequence, PeaGB1 must act as an autoreceptor in this organ. A widespread distribution of PeaGB1 in almost all neuropiles was detected in the cockroach brain. In the male accessory glands mushroom gland and phallic (conglobate) gland, an intense PeaGB1-like immunoreactivity was measured.

The olive fruit fly, Bactrocera oleae, (Rossi) (Diptera:Tephritidae), is a key pest of olive crops. The sterile insect technique (SIT) is an environmentally benign and species-specific method of pest control, aiming to reduce the reproductive potential of a wild population through the mass-release of sterile insects. Previous olive fly SIT trials, involving the release of gamma-ray sterilised mixed-sex populations, achieved limited success. Key problems included altered diurnal mating rhythms of the laboratory-reared insects, leading to assortative mating between released and wild populations, and low competitiveness of the radiation sterilised mass-reared flies. Consequently, the production of competitive, male-only release cohorts is seen as essential. The RIDL (Release of Insects carrying a Dominant Lethal) system is a transgene-based derivative of SIT, one version of which involves the mass release of insects carrying a female specific lethal transgene (fsRIDL). This thesis describes: 1) the development of fsRIDL olive fly strains and the molecular analysis of transgene insertion and function; 2) the analysis of strain life-history parameters; 3) studies into sexual selection and mating compatibility; 4) a caged proof-of-principle population suppression trial; and, 5) selection dynamics on the fsRIDL trait in caged populations. Olive fly fsRIDL strains were developed with full female-lethal penetrance and repressibility. The lead strain displayed similar life-history and sexual competitiveness traits to those of the wild-type strain from which they were derived. In addition, transgenic males showed photoperiod compatibility and strong sexual competitiveness with field-collected wild olive flies. The feasibility of the fsRIDL approach was demonstrated when repeated male releases caused eradication of caged olive fly populations. Although needing field confirmation, these results suggest that fsRIDL olive fly strains may help to mitigate key problems experienced in previous olive fly SIT trials, and could help form the basis of a renewed effort towards olive fly SIT control.

Tanzania’s Ruaha landscape is an international priority area for large carnivore conservation, harbouring roughly 10% of the world’s lions, and important populations of leopards and spotted hyaenas. However, these large carnivore populations are threatened by intense retaliatory killing due to human-carnivore conflict on village land around Ruaha National Park (RNP), mostly as a result of livestock predation by lions, leopards and spotted hyaenas. Moreover, a current lack of ecological data on the distribution of these carnivores hinders the development of effective strategies for conservation and targeted conflict mitigation in this landscape. This study aimed to identify the most significant ecogeographical variables (EGVs) influencing the distribution of lions, leopards and spotted hyaenas across the Ruaha landscape, and to map areas of conservation importance for these species. In addition, the study assessed the influence of EGVs on livestock predation risk by these carnivores in the village land around RNP, and generated a predictive map of predation risk. The relative importance of livestock husbandry practices and EGVs in terms of influencing predation risk within enclosures was also investigated. Proximity to rivers was the most important variable influencing the distribution of large carnivores in Ruaha, and contributed to predation risk of grazing livestock. The traditional livestock husbandry adopted in bomas appeared insufficient to alleviate the inherent risk of predation by large carnivores. The study produced the first detailed maps of lion, leopard and spotted hyaena distribution in the critically important Ruaha landscape, and identified likely livestock depredation hotspots. These results will target conflict mitigation approaches around Ruaha, by identifying particularly high-risk areas for livestock enclosures and grazing stock. Improving husbandry in these areas could help reduce livestock depredation and retaliatory carnivore killing, therefore reducing one of the most significant conservation threats in this critically important landscape.

This thesis examines the evolutionary biology of the RNA viruses, a diverse group of pathogens that cause significant diseases. The focus of this work is the relationship between the processes driving the evolution of virus populations within individual hosts and at the epidemic level. First, Chapter One reviews the basic biology of RNA viruses, the current state of knowledge in relevant topics of evolutionary virology, and the principles that underlie the most commonly used methods in this thesis. In Chapter Two, I develop and test a novel framework to estimate the significance of phylogeny-trait association in viral phylogenies. The method incorporates phylogenetic uncertainty through the use of posterior sets of trees (PST) produced in Bayesian MCMC analyses. In Chapter Three, I conduct a comprehensive analysis of the substitution rate of hepatitis C virus (HCV) in within- and between-host data sets using a relaxed molecular clock. I find that within-host substitution rates are more rapid than previously appreciated, that heterotachy is rife in within-host data sets, and that selection is likely to be a primary driver. In Chapter Four I apply the techniques developed in Chapter Two to successfully detect compartmentalization between peripheral blood and cervical tissues in a large data set of human immunodeficiency virus (HIV) patients. I propose that compartmentalization in the cervix is maintained by selection. I extend the framework developed in Chapter Two in Chapter Five and explore the Type II error of the statistics used. In Chapter Six I review the findings of this thesis and conclude with a general discussion of the relationship between within- and among-host evolution in viruses, and some of the limitations of current techniques.

Biologists have long used demographic approaches to answer questions in ecology and evolution. The utility of these approaches has meant a constant development and refinement of methods. A key milestone has been the development of phenotype structured population models that link ecology and evolution. Moreover, biostatistical research steadily improves methods to coax demographic information from scarce data. In this thesis, I build upon some of the recent advances in the field. My first three studies focus on the consequences of sex differences in life history for population dynamics. Firstly, I test whether males matter for the dynamics of African lion (Panthera leo) populations via a previously unquantified mechanism: the inheritance of phenotype from father to offspring. Secondly, I develop a method to estimate age-specific mortality rates for both sexes in species where one of the sexes disperses around the age of maturity. Thirdly, I apply this method to study variation in mortality between the sexes and between two populations of African lions. After these three chapters, which make contributions to the field of sex-structured population dynamics, I focus on the integration of phenotype structured modelling and quantitative genetics. I illustrate how heritability of a quantitative character that develops with age depends on (i) viability selection, (ii) fertility selection, (iii) the development of the phenotype with age, and (iv) phenotype inheritance from parents to offspring. Our results question the adequacy of quantitative genetics methods to obtain unbiased estimates of heritability for wild populations. This thesis advances our understanding of population development over ecological time scales. This knowledge has applications in conservation and population management, but also contributes to untangling evolutionary processes in wild animals.

Sex chromosomes experience distinct evolutionary environments, due to their unusual pattern of inheritance, and studies of sex chromosome evolution can shed light on the fundamental evolutionary forces acting across the genome as a whole. Here, I combine genomic and transcriptomic data across a wide range of avian species to explore the evolutionary processes governing sex chromosome evolution. Birds are female heterogametic and therefore it is possible, via comparisons with male heterogametic species, to identify the fundamental factors driving sex chromosome evolution, versus those associated with sex. In this thesis, I uncover a complex mosaic of recombination suppression between the Z and W chromosomes, characterized by repeated and independent divergence of gametologs, together with ongoing genetic exchange. Additionally, I highlight the role of mating system, and interplay between evolutionary forces, in driving coding and expression evolution on the Z and W chromosomes. My findings indicate that although the Z chromosome is masculinized for male-specific effects, the magnitude of genetic drift acting on Z-linked genes is elevated in promiscuous relative to monogamous mating systems. In contrast, evolution of the female-limited W chromosome is governed predominately by purifying selection. Together, my results suggest that the role of the Z chromosome in encoding sexual dimorphisms may be limited, but that W-linked genes play a significant role in female-specific fitness. In conclusion, my findings reveal the power of mating system in shaping broad patterns of genome evolution.

Semi-aquatic mammals have evolved to forage in both aquatic and terrestrial environments, and they generally lack locomotor specialisation for either. Having relatively unspecialised adaptations, some semi-aquatic species are generalist foragers, and their activity and diving patterns provide insight into constraints on their foraging behaviour. The recent miniaturisation and improved accuracy of logging devices allow remote data collection from small (<1kg), shallow-diving species. The development of analytical methods has lagged behind technological advances (and techniques developed for fully aquatic animals do not account for the variability of behaviour typical of semi-aquatic animals and their switching between terrestrial and aquatic environments). I applied and developed novel analytical techniques to identify activities and diving patterns of a semi-aquatic mustelid, the American mink (Neovison vison), fitted with Time-Depth Recorders (TDRs). Using a hidden Markov model (HMM) algorithm allowed me to classify dives into three states to identify clustering, and describe sequential diving patterns by mink. TDRs were used to determine active/inactive periods on the basis of rapid TDR temperature changes, and this method was validated empirically. Having developed this methodology, I applied it to 18 datasets collected from 14 mink in lowland England. Terrestrial activity of mink was positively related to ambient temperature (across both sexes), however aquatic activity, especially diving, appeared to be more influenced by daylight than by temperature. Mink showed intersexual differences, with males being more nocturnal and more active on land, and females more diurnal, and more persistent in diving. There was considerable variability between sexes and individuals. This is the first study to use HMM to classify the dives of a semi-aquatic animal, and the first to use TDR temperature records to identify mammalian activity patterns. These methods will be generally applicable to animals that make rapid transitions between environments and have thus far been difficult to study.

Insects are often infected with facultative endosymbiotic bacteria, which can have a range of important ecological effects. The grain aphid, Sitobion avenae, harbours diverse facultative symbionts, which suggests their importance in grain aphid biology. This thesis attempts to explain the ecological roles of the facultative endosymbionts in S. avenae. It also examines the question of whether the horizontal transmission of symbionts between aphid clones and species can be important for shaping the ecology and evolution of multi-species aphid communities. Novel techniques developed for research with the grain aphid study system are presented. Grain aphid clones vary in their tolerance to low temperatures, but this trait is not affected by their facultative endosymbionts. Strains of a symbiont Hamiltonella defensa do not protect grain aphids from hymenopterous parasitoids, regardless of the host genotype. However, experienced parasitoid females preferentially oviposit in aphids which do not harbour symbionts. Comparison of the fitness consequences of infection with the same Hamiltonella strains in their original and in novel grain aphid host clones reveal no consistent differences. Symbiont strains establish easily following artificial transfer between clones of the grain aphid, but the symbionts transferred from other aphid species form less stable infections. Hamiltonella strains do not affect the fecundity of their grain aphid host clones regardless of their host species of origin, but also do not generally confer protection against parasitoids. There are no clear patterns in the distribution of parasitoid-resistant phenotypes across phylogenetic trees of Hamiltonella and its bacteriophage APSE. Strains of four unrelated species of endosymbionts, Rickettsia, Spiroplasma, Rickettsiella and Regiella, confer the same pathogen-resistant phenotype to a single pea aphid clone. The same symbiont strains can confer resistance to clones of two different aphid species. Some strains in multiple infections may compensate for the costs of infections with other symbionts. The importance of these results for understanding the ecological and evolutionary role of facultative endosymbionts in aphids and other insects are discussed, and directions for further research are proposed.

This thesis centres around two genera of large peltospirid gastropods (Mollusca: Neomphalina: Peltospiridae) endemic to hydrothermal vent ecosystems. One is the 'scaly-foot gastropod', an emblematic species of the Indian Ocean vents with unique dermal sclerites covering the foot like roof tiles. The other was recently discovered from expeditions to the Southern and Indian oceans, lacks sclerites and possesses large opercula. As both genera and their assigned species remained undescribed, they were formally described herein which forms a basis to understanding their biology. The 'scaly-foot gastropod' from both the Central Indian Ridge (CIR) and the Southwest Indian Ridge (SWIR) were confirmed to represent a single species and is formally named as Chrysomallon squamiferum. Through molecular genetic analyses using the COI gene, genetic differentiation between SWIR and CIR populations was detected for the 'scaly-foot gastropod'. In contrast, the peltospirids with large opercula from the East Scotia Ridge (ESR) and the SWIR proved to be two distinct species within an undescribed genus. The ESR species was formally described as Gigantopelta chessoia and the SWIR species as G. aegis. The molecular genetic analyses of the COI gene, confirmed the genetic isolation of the two and consolidated their status as separate species. A 3D tomographic model of Chrysomallon squamiferum was generated to characterise the soft anatomy and morphology as well as to understand its internal anatomy and adaptation which remained little-studied. Further to the enlarged esophageal gland already known to house chemosynthetic endosymbionts, C. squamiferum was discovered to have a hypertrophied circulatory system with a gigantic, muscular heart and large ctenidium to adapt to life in a hypoxic environment and to supply the endosymbionts with necessary chemicals. Histological examinations of the sclerites and operculum showed that it was unlikely that the sclerites originated from operculum duplication. Comparisons with polyplacophoran scales revealed starkly different secretion mechanisms despite the superficial similarity, which has implications on the placement of sclerite-bearing Cambrian taxa. Overall, the results from this thesis ascertained the systematic positions of these large-sized, enigmatic peltospirids, and led to improved understanding of their ecology and evolution. The important role of larval dispersal in maintaining metapopulations across the distribution of a vent-endemic taxa is highlighted. The adaptations of vent-endemic taxa remains little-known even in well-studied species, warranting future studies on these and other species.