Thesis (doctoral)--Ludwig-Maximilians-Universität München, 2005. / Title from PDF t.p. (viewed on May 10, 2006). Vita. Includes bibliographical references.

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There is ample evidence that climate change is impacting on phenology and it has been suggested that this may generate trophic mismatches. A key system for investigating phenology and trophic mismatch occurs in spring in temperate deciduous woodlands, where folivorous caterpillars and their predators, insectivorous passerines, are reliant upon ephemeral resources for reproductive success and survival. However, studies are primarily conducted within single-site, oak- (Quercus sp) dominated woodland and focus on a single caterpillar species, winter moth (Operophtera brumata), despite these passerines being habitat generalists with large geographic ranges. It remains to be seen whether insights gained from these studies can be generalised on the landscape scale across different habitats. In this thesis, I explore the extent to which geographic and habitat variation operates in this system and attempt to expand the system beyond a linear single-species food chain into a more biologically realistic multi-species food web. I also identify the most important environmental factors predicting the phenology of the passerines to allow better predictions of how their phenology could alter under future climate change scenarios. To address these questions, I established a novel 220km transect of Scotland incorporating 40 field sites that vary in elevation and the type of deciduous woodland habitat, monitoring six blue tit (Cyanistes caeruleus) nestboxes, tree and invertebrate phenology and abundance, at each site throughout the springs of 2014-16. Firstly, I assess how blue tit occupancy and productivity are affected by the variation in fine-scale woodland habitat, latitude, elevation and prey availability that exists along the transect (Chapter 2). I find that habitat variables strongly affect fledging success but not occupancy or clutch size, whilst occupancy exhibits biogeographic trends, revealing that the relationship between breeding decisions and outcomes differs among habitats and implies that it may be difficult to generalise results from one habitat to others. Next, I aim to identify the environmental aspects which play a role in regulating blue tit reproductive phenology by examining the ability of temperature, tree phenology, invertebrate prey abundance and photoperiod to predict nest initiation and laying dates (Chapter 3). I find that night-time temperature in early spring is the most important predictor of both nest initiation and lay date (slopes ~ -3days/°C) and I suggest that this supports the hypothesis that temperature acts as a constraint on timing rather than a cue. Invertebrate abundance is also a positive correlate of lay date, possibly allowing fine-tuning of timing. This knowledge provides clearer foundations from which to predict future phenological change and possible trophic mismatch in this system. There is the potential that the apparent effect of temperature on blue tit reproductive phenology is indirect and mediated by diet, which is largely undescribed in the period prior to breeding. Therefore, in Chapter 4 I examine how blue tit diet varies across habitat, geography and time, and whether there is a dietary cue utilised to initiate breeding phenology, using data from metabarcoding faeces collected from nestbox-roosting adults in early spring. Geographic variation in diet is substantial, with high site-to-site dietary turnover (β-diversity), as well as high turnover along the elevational and latitudinal gradients studied. Dietary α-diversity (richness) is unaffected by geographical variables, but increases over time, with significant pre-breeding dietary increases in Lepidoptera and Hemiptera signifying a possible cue. In addition, these data provide the most comprehensive next-generation insights into the diet of a wild bird to date and identify 432 prey taxa. Finally, I analyse how biogeographic and habitat variables affect the phenology, abundance and diversity of caterpillars (Chapter 5). Host tree species’ varied significantly in their likelihood of hosting a caterpillar, with oak and willow (Salix sp.) the most likely. Biogeography had less effect on the likelihood of caterpillar occurrence, but elevation delayed peak date by 3.7 days/100m increase. There was also support for the spring caterpillar peak being dominated by a few key species, with over half of all caterpillars identified being of just three of the 62 total species, including winter moth. These findings contribute to understanding how the temporal distribution of caterpillars varies across habitats on the landscape scale. Taken together, the findings of this thesis reveal considerable geographic and habitat variation throughout this system, in both the composition of the food web and the impacts on blue tit productivity, demonstrating why caution must be exercised when extrapolating findings from one location or habitat to others.

Great tits (Aves: Passeriformes; Parus major) and blue tits (Parus caeruleus) nested in boxes in Wytham Woods near Oxford. The breeding densities of both species were limited by the availability of nest sites. The larger great tits were dominant in obtaining nest boxes. This was most important where breeding sites were scarce. The two species did not maintain interspecific territories or interfere with interspecific nest site spacing beyond the immediate vicinity of the nest. Blue and great tit numbers fluctuated in parallel where nest sites were not limiting resources. Annual changes in breeding numbers of great tits were negatively related to blue tit breeding density but great tit density did not seem to affect changes in the blue tit population. Overlap in the feeding sites of blue and great tits was greatest during the summer and interference competition was lowest at this time. The nestling diets of the two species were very similar. Despite an apparent abundance of food for nestlings, adults were pressed to feed large broods. Food for nestlings was probably a limiting and depletable resource. The date of clutch initiation of great tits, but not blue tits, was retarded at high densities of blue or great tits. The clutch sizes of both species were probably negatively affected by high breeding densities of congeners but the results were not clear cut. Heavy great tit fledglings are most likely to survive to breed. Great tit fledglings were heavier at low densities of blue tits. An experiment in which blue tit young were removed from a section of the Woods, showed that great tit nestlings were heavier and developed faster, and that female condition was better, than in a control section or section where blue tit broods were supplemented. In terms of resource competition, blue tits were the better scramble competitors and great tits the better interference competitors.

Depuis plusieurs décennies, les populations font face à des changements environnementaux sans précédent. Afin de répondre à ces changements les espèces peuvent se déplacer vers un autre endroit (c’est la dispersion), répondre aux changements par plasticité phénotypique, ou encore s’adapter par sélection naturelle (c’est la réponse évolutive). Si la dispersion et la plasticité phénotypique permettent une réponse rapide face à un changement environnemental tel que le réchauffement climatique, seule la réponse évolutive permet une adaptation durable. Une réponse évolutive au changement climatique peut être attendue seulement si l'augmentation des températures occasionne une pression de sélection sur un et / ou des traits. Cependant, quantifier les pressions de sélection qui agissent sur les populations naturelles reste difficile, en particulier celles qui accompagnent le changement climatique.Au cours de cette thèse, j'ai cherché à quantifier les pressions de sélection reliées au changement climatique agissant sur une population sauvage de mésange bleue (Cyanistes caeruleus) suivie depuis 26 ans près de Montpellier. Dans un premier temps, j'ai exploré le biais induit par l'autocorrélation spatiale sur l'estimation de la sélection naturelle lorsqu'elle n'est pas prise en compte dans une population sauvage. Dans un deuxième temps, j'ai quantifier l'impact du réchauffement climatique sur la force de sélection agissant sur la date de ponte, un trait fortement relié au succès reproducteur chez la mésange bleue. Cependant, si le changement climatique est caractérisé par une augmentation des températures, il s'accompagne aussi d'une augmentation de la fréquence d'évènements extrêmes climatique (EEC). Dans un troisième temps, je me suis donc attaché à estimer l'influence des températures et des pluies extrêmes sur la force de la sélection naturelle. Concernant l'autocorrélation spatiale, nos résultats ont permit de mettre en avant un biais majeur sur l'estimation de la sélection naturelle lorsqu'elle n'est pas prise en compte. En effet, lorsque la valeur sélective est positivement spatialement autocorrélée, un modèle de sélection non-spatialisé surestimera la force de la sélection naturelle agissant sur les traits. Nous avons donc développé 4 modèles de sélection spatialisés, et nous les avons comparé entre eux sur la base de plusieurs paramètres. De plus, nous avons détecté un fort réchauffement dans notre population qui s'accompagnait d'une augmentation de la force de sélection naturelle agissant sur la date de ponte. En effet, nous avons quantifié une augmentation de 46% de la force de sélection naturelle tout les +1°C de la température maximum du mois d'Avril. Au delà des températures moyennes, nous avons mis en évidence un fort impact négatif de la présence / absence de journées extrêmement chaudes durant l'élevage des oisillons au nid sur le succès d'envol. Plus intéressant encore, ces EECs augmentaient significativement la force de sélection naturelle agissant sur la date de ponte, indépendamment de l'augmentation des températures moyennes.Ainsi, ma thèse a contribué à une meilleure compréhension méthodologique de la sélection naturelle, ainsi que de l'impact sélectif du changement climatique. Le premier chapitre a permit de mettre en évidence un problème méthodologique majeur lorsque on estime la sélection naturelle dans les populations sauvages, ainsi que de proposer des solutions concrètes. Le second chapitre a permit de confirmer l'impact sélectif du changement climatique, laissant entrevoir une possibilité de réponse évolutive au réchauffement. Enfin, le troisième chapitre démontrait la sélection naturelle reliée à l'apparition d'EEC, et en quantifiait les effets, un résultats encore jamais montré. Nous espérons que cette thèse ouvrira la voie à plus d'études explorant l'impact du changement climatique sur la sélection naturelle, permettant peut être de prévoir les réponses évolutives futures. / Global change results in an increase in temperature mean and variability and generates new environmental conditions for wild populations. In order to respond to this new treat, wild populations can move to another place (dispersion), change their behaviour or their phenology (phenotypic plasticity) or respond by evolutionary change (involving changes in alleles frequency). However, phenotypic plasticity or dispersion are unlikely to sustain population responses to climate change over the long term, contrary to the evolutionary response. Wild populations could evolutionary respond to climate change only if the increase in temperature represents a selective pressure on fitness-related traits. Yet, quantifying the selective pressures acting on wild populations remains challenging, especially in response to climate change.During this PhD, I focused on the selection pressures related to climate warming during a 26-year monitoring of blue tits (Cyanistes caeruleus) in southern France. First, I explored the bias inducted by spatial autocorrelation on estimation of natural selection in the wild when it is not taken into account. Second, I quantified the impact of climate warming on natural selection acting on laying date. Along with a rise in mean temperatures, climate change is also characterized by an increase in the frequency of extreme climatic events (ECE). Hence, I finally explored the impact of multiple ECEs such heavy rainfalls and extremes temperatures on the strength of natural selection acting on laying date.Our results showed that spatial autocorrelation is severely biasing our estimation of natural selection in the wild if it is not taken into account. In fact, natural selection is continuously overestimated by a non-explicit spatial model when the fitness is spatially autocorrelated. Hence, we developed 4 spatially-explicit models of selection, and we compared them in order to assess their reliability. Also, we detected a strong warming in our study site, which was correlated with an increase in natural selection strength. In fact, our results indicated an increase in the strength of selection by 46% every +1°C anomalies in maximum April temperature. Beyond this selective impact of the warming trend, we detected an impact of the extreme hot days occurrence during the nestlings stage on the fledglings success. More interesting, these ECEs significantly increased the strength of selection acting on laying date, independently form the mean temperatures.To conclude, my PhD helped to a better understanding of natural selection estimation in the wild, and the selective impact of climate warming. The first part allowed to develop specific models of selection in order to take into account spatial autocorrelation, a universal phenomenon present at any scale. The second part confirmed the selective impact of warming, pointing out a potential evolutionary response to climate change in the future. The last part demonstrated and quantified the selective impact of multiple ECEs, which has not been demonstrated before. We call now for more studies assessing the selective impact of climate warming, which could allow to predict the evolutionary responses in the future.

In nature, environments are often variable and heterogeneous influencing ecological and evolutionary processes. This thesis focus on how animals interact with their environment and how that affects the reproductive decisions they make. Using empirical data collected from wild collared flycatcher populations, experiments and molecular approaches I try to unveil some of these relationships and the evolutionary, ecological and conservation implications of these findings are discussed. Firstly, collared flycatchers were shown to use breeding densities of their own and other species using similar resources when assessing costs and benefits associated to breeding in specific habitats. However, species will vary in how informative they are, and the worst competitor – with whom you overlap most in resources needs – also provides the best source of information. Collared flycatcher parents will also benefit differentially from investments in sons and daughters due to habitat characteristics and dispersal differences between the sexes. Here, I show that they will produce more of the sex that will give the highest expected fitness return given the environment they are in. These results also provide a reciprocal scenario to Clark's (1978) classical study of sex ratio adjustment in relation to local resource competition (LRC), as more of the natal philopatric sex is produced when LRC is low. Secondly, the effect of elaborated ornaments on paternity in the socially monogamous collared flycatcher was shown to be of more importance in areas where the intensity of intra- and intersexual conflicts are expected to be elevated. Hence, ornamentation by environmental interactions determines paternity, illustrating that sexual selection through extra-pair paternity is context dependent. Finally, even though the collared flycatcher populations that this thesis is based on have been studied on their breeding grounds for more then 25 years, we know little of where they are when they are not breeding. Here, stable isotope signatures in winter-grown feathers suggests that they may spend their winter with their breeding ground neighbours and do so repeatedly over years. Differences between breeding populations at this small scale should have many impactions for evolutionary and ecological processes as it will, for example, determine with whom individuals interact throughout their life.

By warning signalling prey advertises its unpalatability to the predators. Typical examples are true bugs (Heteroptera) combining optical and chemical signals. The aim of this study was to find out the effectiveness of chemical defence against wild-caught adults of great tits (Parus major) and blue tits (Cyanistes caeruleus) and towards hand-reared juveniles of great tits. The tested prey were adults and larvae of two invasive species of genus Oxycarenus (Heteroptera: Oxycarenidae) (aposematic O. lavaterae, non-aposematic O. hyalinipennis), adults of Horvathiolus superbus (Heteroptera: Lygaeidae) and crickets (Gryllus assimilis) as a control prey. We were focusing on the influence of seed bugs to the initial reaction, the learning process and to the displays of discomfort behaviour in tits. Reactions affected by the tit species, age and sex were compared in adults. Juveniles were divided into two independent experimental groups, one group was offered adults of genus Oxycarenus, the second was offered sunflower (Helianthus sp.) or Digitalis sp. seed fed adults of H. superbus. The first bug offered did not elicit an initial aversive reaction in tits, but birds reacted aversively to the bugs after having further handling experience. All juveniles, contrary to the adult tits, attacked at least one bug...

Innerhalb des Zeitraumes 2002 bis 2004 wurden in Berlin Untersuchungen zur Nahrungsökologie verschiedener Vogelarten durchgeführt, wobei auch die Konsequenzen wie der Fortpflanzungserfolg analysiert wurden. Neben der Aufnahme der Brutvogel- (2002) und Wintervogelgemeinschaft (2002/2003) auf fünf Pf à ca. 20 ha entlang eines Urbangradienten, wurden die Arten Blaumeise, Grünling, Haussperling und Turmfalke speziell untersucht. Auf den fünf Pf, die sich hinsichtlich Baustruktur, Flächennutzung und Vegetationsanteil unterschieden, wurden 35 Brutvogelarten festgestellt. Anhand der Artenzahlen ist deutlich ein Urbangradient zu erkennen: Das Stadtzentrum (Ze) wies 12, das Gewerbegebiet (Ge) 15, die Hochhaussiedlung (Ho) 17, die Einfamilienhaussiedlung (Eh) 18 und der Park (Pa) 28 Arten auf. Im Verlauf der Wintervogelkartierung wurden insgesamt 3763 Individuen in 30 Arten registriert. Im Pa wurde bei der Blaumeise der höchste Reproduktionserfolg mit knapp 5,9 Ausgeflogenen/Brut festgestellt. Anhand des Zustandes der Nestlinge und des Fortpflanzungserfolgs sowie durch die Dokumentation des Fütterungsverhaltens der Altvögel anhand von Camcorderaufnahmen wurde deutlich, dass das Nahrungsangebot für die Meisen im Pa am besten war. Der carnivore Turmfalke wurde 2002 bis 2004 unabhängig von den fünf Pf an zehn Nistkastenstandorten untersucht, die in drei unterschiedlichen Zonen lagen: City (Ci), Mischgebiet (Mi) und ländliche Zone (LZ). Als Besonderheit wurden in jedem Nistkasten in der Ci in jedem Jahr anthropogene Nahrungsreste, z. B. Kotelettknochen, gefunden, was zeigt, dass sich der Turmfalke, neben der Nutzung von Gebäuden als Nistplatz, auch hinsichtlich seiner Nahrungsökologie an den Menschen anpasst. Dass die Blaumeise, der Grünling und der Haussperling insgesamt in Berlin gut zurechtkommen, selbst in der Innenstadt, indiziert die Beutetierliste des Turmfalken: Der Haussperling stand an erster Stelle, der Grünling an zweiter und die Blaumeise an sechster Stelle der gefangenen Vogelarten. Besonders in einer Metropole wie Berlin besteht weiterer Forschungsbedarf, da die Synurbanisierung verschiedener Vogelarten ständig voranschreitet. / From 2002 to 2004 the feeding ecology of different bird species was studied in Berlin. Thereby consequences like the success in reproduction were also examined. Beside the survey of the breeding bird (2002) and the winter bird community (2002/2003) in five study sites each covering an area of approx. 20 ha along an urban gradient, the species Blue Tit, Greenfinch, House Sparrow and Kestrel were researched in detail. In total, 35 breeding bird species were found in the five sites, which were characterised by different building structure, land use and vegetation cover. Considering the species` number, an urban gradient was clearly visible: In the city centre (Ze) 12 species were recorded, in the industrial area (Ge) 15, in the high-rise apartment building area (Ho) 17, in the detached family housing (Eh) 18 and in the park (Pa) 28 species. During the winter bird census a total of 3,763 individuals and 30 bird species was recorded. In the Pa, the highest reproduction success of the Blue Tit (5,9 fledglings/brood) was recorded. On the basis of the nestlings'' condition and the reproduction success, as well as by documentation of the feeding behaviour of the adult birds by means of video recording (camcorder), it became clear, that food supply was best in the Pa for the Tits. From 2002 to 2004, the carnivorous Kestrel was examined independent from the five study sites at ten nesting sites (nest boxes), which were located in three different zones: City (Ci), Mixed Zone (Mi) and Outskirts (LZ). Striking were anthropogenic food items, like cutlet bones, found in every nest box in the Ci in all three years, which shows the adaptation to humans not only with regard to nesting sites (buildings) but also with regard to feeding ecology. The prey species list of the Kestrel indicates that Blue Tit, Greenfinch and House Sparrow get along well all in all, also in the city centre: the House Sparrow ranked first, the Greenfinch second and the Blue Tit sixth of the captured bird species. Especially in a metropolis like Berlin there is need for further research, because the synurbization of different bird species progresses permanently.

Avian malaria (Plasmodium spp.) is a popular model system to study the ecology and evolution of parasite-host-vector interactions in the wild. These studies have historically focused mostly on the avian hosts and the malaria parasites. Knowledge regarding the role of vectors is essential to our understanding of these wild systems, but has only very recently started to accumulate. This thesis aimed to contribute to this field by assessing mosquito-malaria-host associations for British mosquitoes and the role of mosquito ecology in shaping these parasite systems in a British woodland study site, using molecular, field ecology and statistical modelling methodologies. From the 12 mosquito species or species groups found, I showed that the Cx.pipiens/torrentium mosquito group is likely to have a major role in avian malaria transmission in Great Britain, while Cs. annulata may be transmitting P. circumflexum. I also demonstrated a positive spatial association between mosquito density per host and avian malaria prevalence, in accordance with theoretical expectations for malaria transmission. Findings here provide evidence that avian malaria transmission in British woodlands is limited mainly to June-August, being preceded by relapse of previous infections or, alternatively, by maintenance of chronic blood parasitaemia through the colder months; this agrees with theoretical expectations and findings elsewhere for temperate climates. This thesis also described local-scale spatial heterogeneity and seasonal variation in adult mosquito abundance within a British woodland where avian malaria is endemic, with differing patterns found between species or species groups. Spatially, variation in adult mosquito abundance was associated with microclimatic and landscape variables such as distances to mosquito breeding sites, microclimate and canopy height; seasonally, variation in mosquito abundance was associated with temperature and rainfall, alongside calendar date. The heterogeneity in mosquito parameters and associations with environmental variables found at a site where avian malaria is endemic highlights the need to anticipate such complexity when trying to understand Plasmodium transmission. By doing so, we further extend the potential of these parasite systems to improve our knowledge regarding the ecology and evolution of parasite-host-vector associations.