Altitudinal variation in haemosporidian parasite distribution in great tit populations.

BACKGROUND: One of the major issues concerning disease ecology and conservation is knowledge of the factors that influence the distribution of parasites and consequently disease outbreaks. This study aimed to investigate avian haemosporidian composition and the distribution of these parasites in three altitudinally separated great tit (Parus major) populations in western Switzerland over a three-year period. The objectives were to determine the lineage diversity of parasites occuring across the study populations and to investigate whether altitudinal gradients govern the distribution of haemosporidian parasites by lineage. METHODS: In this study molecular approaches (PCR and sequencing) were used to detect avian blood parasites (Plasmodium sp., Haemoproteus sp. and Leucocytozoon sp.) in populations of adult great tits caught on their nests during three consecutive breeding seasons. RESULTS: High levels of parasite prevalence (88-96%) were found across all of the study populations with no significant altitude effect. Altitude did, however, govern the distribution of parasites belonging to different genera, with Plasmodium parasites being more prevalent at lower altitudes, Leucocytozoon parasites more at high altitude and Haemoproteus parasite prevalence increasing with altitude. A total of 27 haemosporidian parasite lineages were recorded across all study sites, with diversity showing a positive correlation to altitude. Parasites belonging to lineage SGS1 (P. relictum) and PARUS4 and PARUS19 (Leucocytozoon sp.) dominated lower altitudes. SW2 (P. polare) was the second most prevalent lineage of parasite detected overall and these parasites were responsible for 68% of infections at intermediate altitude, but were only documented at this one study site. CONCLUSIONS: Avian haemosporidian parasites are not homogeneously distributed across host populations, but differ by altitude. This difference is most probably brought about by environmental factors influencing vector prevalence and distribution. The high occurrence of co-infection by different genera of parasites might have pronounced effects on host fitness and should consequently be investigated more rigorously.

Ectoparasite affects choice and use of roost sites in the great tit, Parus major

Many diurnal bird species roost at night in holes. As a regular visitor of a hole they are therefore a welcome host for several species of ectoparasites. The interactions of ectoparasites with the behaviour, life-history traits and population demography of their hosts are largely unknown. In the present study the effects of the haematophagous hen flea, Ceratophyllus gallinae , on the great tit's choice of winter roost site were investigated experimentally. Three experiments tested (1) whether great tits prefer a clean nestbox to one containing an old, but parasite-free nest, (2) whether they prefer a parasite-free nestbox to one infested with the haematophagous hen flea, and (3) whether they prefer not to use a nestbox when there is only an infested box available in their territory. In the first experiment there was no discrimination and both kinds of boxes were used equally often. In the second experiment the great tits clearly preferred to roost in the box without ectoparasites. In the third experiment a significantly higher proportion of the infested nestboxes were not used for roosting compared with the parasite-free boxes. Recently the validity of the conclusions drawn from nestbox studies where the naturally occurring detrimental ectoparasites are eliminated by the routine removal of old nests between breeding seasons has been questioned. This study shows that ectoparasites affect host behaviour and therefore lends support to that criticism.

Parental influence on sibling rivalry in great tit, Parus major, nests

Sibling and parente-offspring conflicts arise mainly over the amount and distribution of parental care, especially food. In altricial bird species where the young depend on parents for obtaining food, parents may control sibling competition by the choice of their respective provisioning locations. In great tits, the parents use fixed provisioning positions on the nest rim that are determined early in the breeding cycle and maintained until. edging. The two parents may choose positions that are close to each other, or far apart, and thereby increase or relax the pressure for optimal feeding positioning among nestlings. As an inspiration to this study we previously found that the two parents provide food from closer positions if the nest is infested by ectoparasites. Here, we tested the hypothesis that the parental choice of relative provisioning locations could be strategically used to control nestling competition. We forced parents to feed from either one or two provisioning locations and assessed the induced change in nestling movement, weight gain, and food distribution among siblings. We show that the angular distance between male and female locations influences the level of behavioural competition and affects nestling weight gain and food distribution. It is the first evidence for hole-nesting birds, where it was assumed that the nestling closest to the entrance hole was fed first, that the apparent choice of feeding positions by parents could be a way of controlling sibling competition and thereby also taking partial control over the outcome of parente-offspring conflict. (c) 2007 The Association for the Study of Animal Behaviour. Published by Elsevier Ltd. All rights reserved.

Post-fledging care, philopatry and recruitment in brown thornbills

1. We describe patterns of post-fledging care, dispersal and recruitment in four cohorts of brown thornbills Acanthiza pusilla. We examine what factors influence post-fledging survival and determine how post-hedging care and the timing of dispersal influence the probability of recruitment in this small, pair breeding, Australian passerine. 2. Fledgling thornbills were dependent on their parents for approximately 6 weeks. Male fledglings were more likely than female fledglings to survive until independence. For both sexes, the probability of reaching independence increased as nestling weight increased and was higher for nestlings that fledged later in the season. 3. The timing of dispersal by juvenile thornbills was bimodal. Juveniles either dispersed by the end of the breeding season or remained on their natal territory into the autumn and winter. Juveniles that delayed dispersal were four times more likely to recruit into the local breeding population than juveniles that dispersed early. 4. Delayed dispersal was advantageous because individuals that remained on their natal territory suffered little mortality and tended to disperse only when a local vacancy was available. Consequently, the risk of mortality associated with obtaining a breeding vacancy using this dispersal strategy was low. 5. Males, the more philopatric sex, were far more likely than females to delay dispersal. Despite the apparent advantages of prolonged natal philopatry, however, only 54% of pairs that raised male fledglings to independence had sons that postponed dispersal, and most of these philopatric sons gained vacancies before their parents bred again. Consequently, few sons have the opportunity to help their parents. Constraints on delayed dispersal therefore appear to play a major role in the evolution of pair-breeding in the brown thornbill.

Paternal investment affects prevalence of malaria.

Both reproduction and parasite defense can be costly, and an animal may face a trade-off between investing in offspring or in parasite defense. In contrast to the findings from nonexperimental studies that the poorly reproducing individuals are often the ones with high parasite loads, this life-history view predicts that individuals with high reproductive investment will show high parasite prevalence. Here we provide an experimental confirmation of a positive association between parental investment levels of male great tits Parus major and the prevalence of Plasmodium spp, a hematozoa causing malaria in various bird species. We manipulated brood size, measured feeding effort of both males and females, and assessed the prevalence of the hemoparasite from blood smears. In enlarged broods the males, but not the females, showed significantly higher rates of food provisioning to the chicks, and the rate of malarial infection was found to be more than double in male, but not female, parents of enlarged broods. The findings show that there may be a trade-off between reproductive effort and parasite defense of the host and also suggest a mechanism for the well documented trade-off between current reproductive effort and parental survival.

Potential evidence of parasite avoidance in an avian malarial vector

Epidemiological studies of malaria or other vector-transmitted diseases often consider vectors as passive actors in the complex life cycle of the parasites, assuming that vector populations are homogeneous and vertebrate hosts are equally susceptible to being infected during their lifetime. However, some studies based on both human and rodent malaria systems found that mosquito vectors preferentially selected infected vertebrate hosts. This subject has been scarcely investigated in avian malaria models and even less in wild animals using natural host-parasite associations. We investigated whether the malaria infection status of wild great tits, Parus major, played a role in host selection by the mosquito vector Culex pipiens. Pairs of infected and uninfected birds were tested in a dual-choice olfactometer to assess their attractiveness to the mosquitoes. Plasmodium-infected birds attracted significantly fewer mosquitoes than the uninfected ones, which suggest that avian malaria parasites alter hosts' odours involved in vector orientation. Reaction time of the mosquitoes, that is, the time taken to select a host, and activation of mosquitoes, defined as the proportion of individuals flying towards one of the hosts, were not affected by the bird's infection status. The importance of these behavioural responses for the vector is discussed in light of recent advances in related or similar model systems.

Nestling detectability affects parental feeding preferences in a cavity-nesting bird

In many avian species, nestlings have evolved striking plumage, behaviours and mouth colours to obtain a greater share of parental investment. Studies revealing parental feeding preferences for nestlings with red gapes have proposed that red mouth colour in songbirds can act as a signal of nestling need or condition. Alternative hypotheses suggest that bright nestling mouths in cavity-nesting birds evolved to increase nestling detectability by the parents. We tested whether nestling mouth colour affects parental feeding preferences in great tits, Pants major L. In broods of six young, we experimentally painted mouth gapes and flanges either red or yellow and tested the effect of mouth colour on nestlings' mass gain under two lighting conditions. In nests with high luminosity, there was no significant effect of mouth colour on mass gain. In nests with low luminosity, nestlings with red gapes and flanges gained less mass than nestlings with red gapes and yellow flanges or both yellow gapes and flanges. Our results suggest that, in nests with low luminosity, red mouths decreased nestling detectability to the feeding parents and support the hypothesis that poor luminosity in nesting cavities can select for pale mouths. Overall, our results do not support the hypothesis that red mouth colour signals nestling need or condition to parent great tits.

Characterising avian haemosporidian communities : ecological factors, parasite persistence and co-infection

Les parasites jouent un rôle clef dans l'évolution des comportements et des traits d'histoire de vie de leurs hôtes. Le parasitisme s'avère parfois dévastateur à l'échelle de population d'hôtes, et peut également altérer certains traits associés à la valeur sélective d'un individu infecté, tels que son succès reproducteur ou encore son taux de mortalité. La coévolution hôte/parasite, qui représente l'une des forces sélectives les plus puissantes dans l'évolution des organismes, peut également conduire les partenaires de l'association parasitaire à s'adapter localement à des environnements hétérogènes. Cette thèse porte sur l'étude de parasites aviaires, du genre Plasmodium, Haemopro- teus et Leucocytozoon (Haemosporidae), naturellement associés à différentes populations de mésanges charbonnières (Parus major) et d'hirondelles des fenêtres (Delichon ur- bicum). Dans un premier temps, nous avons cherché à déterminer comment se distribuent ces parasites au sein de différentes populations hôtes et si ces communautés de parasites sont structurées. Par la suite, la principale question à laquelle nous voulions répondre était de savoir comment ces parasites, et notamment après coexistence de plusieurs lignées génétiques d'Haemosporidae au sein dun même-individu (i.e. co-infection), affectent la physiologie et le succès de reproducteur des hôtes. Nos résultats suggèrent que la distribution des Haemosporidae est principalement gouvernée par la présence d'insectes vecteurs et que la persistance de l'infection chez les hôtes varie en fonction du genre d'Haemosporidae (Chapitre 1-2). Par ailleurs, nous avons trouvé que des lignées de parasite génétiquement distinctes peuvent avoir des effets contrastés sur leurs hôtes. Par exemple, les hôtes exhibent des différences de parasitémie marquées en fonction des lignées de parasites responsable de l'infection. De plus, le succès reproducteur ainsi que la charge parasitaire des mésanges infectées par Plasmodium ou Haemoproteus n'étaient pas affecté par l'infection simultanée avec Leucocytozoon (Chapitre 2-3). Dans le Chapitre 4, j'ai examiné la capacité immunitaire de mésanges charbonnières infectées par des hémosporidies. Les résultats n'ont pas été concluant, et je suggère fortement une réévaluation de ceux-ci dans de futures études. Les mésanges charbonnières ne semblent pas signaler leur statut infectieux par la coloration de leur plumage (Chapitre 5); toutefois, la coloration noire des plumes reflète l'état de stress oxydatif des mésanges, qui dépend lui-même de l'infection parasitaire. La coloration verte pourrait également indiquer la qualité des soins paxentaux délivrés par les mésanges adultes femelles à leurs petits, comme le suggère la corrélation que nous avons observée entre la masse des jeunes d'une nichée et la coloration de leur mère. Les hirondelles capturées en Algérie souffrent plus de l'infection que celles échantillon¬nées en Europe (Chapitre 6). Les similitudes observées entre les communautés de par¬asites affectant les populations européennes et celles des populations nord-africaines suggèrent que la transmission des parasites a lieu lors de la migration vers le sud. A l'instar de nos observations sur les mésanges dans les chapitres 2 et 3, les hirondelles co-infectées ne montrent pas d'altérations de leur condition physique. Cette thèse démontre qu'il existe, au sein des populations de mésanges charbonnières, des interactions antagonistes entre, d'une part, les parasites et leurs hôtes et d'autre part, entre différent parasites. Le résultat de ces interactions antagonistes varie en fonction des espèces et de la zone géographique considérée. Nous avons démontré que les interactions ne suivent pas toujours la théorie, puisque la coevolution qui, en suivant le concept de la virulence, devrait augmenter la charge parasitaire et diminuer la condition physique des hôtes, ne montre pourtant pas d'impact négatif sur les populations de mésanges. Nous pouvons maintenant concentrer nos efforts à la caractérisation des interactions antagonistes. De plus, grâce aux avancées des méthodes moléculaires, nous pouvons suivre et étudier en détails comment ces interactions se manifestent et quels sont leurs effets sur la condition physique des hôtes. - Parasites are key in shaping various behavioural and life-history traits of their hosts. The influence of parasitism on host populations varies from slight to devastating and might influence such parameters as mortality rates or reproductive success. Host-parasite coevolution is one of the most powerful selective forces in evolution and can lead to local adaptation of parasites and hosts in spatially structured environments. In this thesis, I studied haemosporidian parasites in different populations of great tits (Parus major) and house martins (Delichon urbicum). Firstly, I wanted to determine how parasites are distributed and if parasite communities are structured. The main question I wanted to address hereafter was how parasites, and specifically infection with multiple genera of parasites (i.e. co-infection) influenced host physiology and reproductive success. I found that parasite distribution is environmentally driven and could therefore be closely linked to vector prevalence; and that the stability of parasite infection over time is genus-dependent (Chapter 1 - 2). I further found that different haemosporidian lineages might interact differently with their hosts as parasitaemia was strongly lineage-specific and that the presence of Leucocytozoon parasites showed no correlation to Plasmodium or Haemoproteus parasitaemia, nor to great tit reproductive success (Chapter 2-3). In Chapter 4 I examined immune capacity of haemosporidian-infected great tits. The results proved inconclusive, and I strongly suggest re-evaluation hereof in future work. Great tits do not appear to signal parasite infection through plumage colouration (Chapter 5); however, infection did have a link to oxidative stress resistance which is strongly signalled through the black breast stripe, with darker males being more resistant and darker females less resistant. Females might incur different costs associated with darker stripes. This would allow reversal of signaling function. Green colouration could also serve as a cue for female provisioning quality as indicated by the strong correlation between colouration and chick body mass. Breeding house martins caught in Algeria suffer greater haemosporidian infection than European populations (Chapter 6). Similar parasite communities in European and North-African populations suggest transmission of parasites may occur during southward migration. Similarly to what was observed in great tits in Chapter 2 and 3, no relationship was found between parasite co-infection and Swiss house martin body condition. This thesis demonstrates that host-parasite and inter-parasite antagonistic interac¬tions exist in great tit populations. How these interactions play out is species dependent and varies geographically. I have demonstrated that interactions do not always follow the theory, as co-infection - which under the concept of virulence should increase parasitaemia and decrease body condition - showed no negative impact on great tit populations. We can now concentrate our efforts on characterising these antagonistic interactions, and with the advance in molecular methods, track and investigate how these interactions play out and what the effect on host fitness is.

Twofold cost of reproduction: an increase in parental effort leads to higher malarial parasitaemia and to a decrease in resistance to oxidative stress.

Parental effort is usually associated with high metabolism that could lead to an increase in the production of reactive oxidative species giving rise to oxidative stress. Since many antioxidants involved in the resistance to oxidative stress can also enhance immune function, an increase in parental effort may diminish the level of antioxidants otherwise involved in parasite resistance. In the present study, we performed brood size manipulation in a population of great tits (Parus major) to create different levels of parental effort. We measured resistance to oxidative stress and used a newly developed quantitative PCR assay to quantify malarial parasitaemia. We found that males with an enlarged brood had significantly higher level of malarial parasites and lower red blood cell resistance to free radicals than males rearing control and reduced broods. Brood size manipulation did not affect female parasitaemia, although females with an enlarged brood had lower red blood cell resistance than females with control and reduced broods. However, for both sexes, there was no relationship between the level of parasitaemia and resistance to oxidative stress, suggesting a twofold cost of reproduction. Our results thus suggest the presence of two proximate and independent mechanisms for the well-documented trade-off between current reproductive effort and parental survival.

The role of vectors in the epidemiology of avian malaria

Studies on host-parasite relationships have commonly reported that parasitized hosts undergo changes in their behavioural and life history traits. How do these changes affect the fitness of the hosts? What are the ecological and evolutionary drivers of these changes? These open questions are crucial to predict the parasite spread amongst hosts. Surprisingly, mosquito vectors of diseases to humans and animals have long been seen as passive parasite transporters, being unaffected by the infection though they also function as hosts. Natural parasite-vector interactions are therefore poorly documented in the literature. In this thesis, we seek to address the role of wild vectors in the epidemiology of avian Plasmodium, the etiological agents of malaria in birds. We first conducted avian malaria surveys in field-caught mosquitoes to identify the natural vectors in our temperate study area. We report that ornithophilic Culex pipiens primarily act as a vector for Plasmodium vaughani in spring, this parasite species being progressively replaced by P. relictum along with the season. Season-related factors may thus shape the mosquitoes' vectorial capacity. We then used experimental approaches to determine the effect of avian malaria on wild, naturally infected C. pipiens. We show that infected mosquitoes incur unavoidable physiological costs associated with parasite exploitation, these costs being expressed as a reduced survival under nutritionally stressed conditions only. These results are of significant importance for the epidemiology of avian malaria since seasonal changes in climate may likely influence food quality and quantity available to the mosquitoes. The host-selection preferences of the vectors with respect to the malaria-infection status of their bird hosts largely determine the disease spreading. In a second laboratory experiment, we thus offered wild C. pipiens the opportunity to choose between uninfected and naturally infected great tits, Parus major. We show that host-seeking mosquitoes have innate orientation preferences for uninfected birds. This suggests that avian malaria parasites exert strong selective pressures on their vectors, pushing them to evolve anti-parasite behaviours. We lastly investigated the links between malaria-associated symptoms in birds and resulting attractiveness to the mosquitoes. We show that experimentally malaria-infected canaries, Serinus canaria, suffer severe haematocrit reduction at peak parasitaemia and reduced basal metabolic rate later in the course of the infection. However, no links between infection and bird attractiveness to the mosquitoes were shown in an experiment using canaries as live bait for mosquito trap in the field. These links may have been masked by confounding environmental factors. Using a system where the vectors, parasites and hosts co-occur in sympatry, this thesis illustrates that vectors are not always Plasmodium permissive, which opposes to the traditional view that malaria parasites should have little effect on their vectors. The way that the vectors respond to the parasite threat is largely determined by the environmental conditions. This may have major implications for the epidemiology of avian malaria. - Les études portant sur les relations hôtes-parasites mentionnent souvent que les hôtes parasités subissent des modifications de leurs traits d'histoire de vie ou bien comportementaux. Comment ces changements affectent-ils la valeur sélective des hôtes et celle de leurs parasites ? Quels sont les déterminants de ces modifications ? Ces questions sont d'un grand intérêt en épidémiologie. Pour autant, les moustiques vecteurs de maladies infectieuses ont longtemps été perçus comme de simples transporteurs de parasites, n'étant pas affectés par ces derniers. Cette thèse porte sur le rôle des vecteurs dans l'épidémiologie des Plasmodium aviaires, agents étiologiques de la malaria chez les oiseaux. Dans le but d'identifier les vecteurs naturels de malaria aviaire dans notre zone d'étude, nous avons tout d'abord collecté des moustiques sur le terrain, puis déterminé leur statut infectieux. Nous rapportons que les moustiques Culex pipiens sont principalement impliqués dans la transmission de Plasmodium vaughani au printemps, cette espèce de parasite étant progressivement remplacée par P. relictum au fil de la saison de transmission. Nous avons ensuite conduit une expérience visant à déterminer l'effet de la malaria aviaire sur des C. pipiens sauvages, naturellement infectés. Nous montrons que des coûts sont associés à l'infection pour les moustiques. Ces coûts occasionnent une diminution de la survie des vecteurs seulement lorsque ceux-ci sont privés de ressources nutritionnelles. Des changements saisonniers de climats pourraient affecter la quantité et la qualité des ressources disponibles pour les vecteurs et donc, leur aptitude à transmettre l'infection. Les traits comportementaux des moustiques vecteurs, tels que la recherche et le choix d'un hôte pour se nourrir, sont d'une importance majeure pour la dispersion de la malaria. Pour cela, nous avons offert à des C. pipiens sauvages l'opportunité de choisir simultanément entre une mésange charbonnière (Parus major) saine et une autre naturellement infectée. Nous montrons que les moustiques s'orientent préférentiellement vers des mésanges saines. Les Plasmodium aviaires exerceraient donc de fortes pressions de sélection sur leurs vecteurs, favorisant ainsi l'évolution de comportements d'évitement des parasites. Enfin nous avons cherché à identifier de potentiels liens entre symptômes de l'infection malarique chez les oiseaux et attractivité de ces derniers pour les moustiques. Nous montrons que des canaris (Serinus canaria) expérimentalement infectés sont fortement anémiés au moment du pic infectieux et que leur métabolisme basai diminue plus tard au cours de l'infection. Toutefois, aucun lien entre le statut infectieux et l'attractivité des canaris pour les moustiques n'a pu être montré lors d'une expérience réalisée en nature. Il se peut que ces liens aient été masqués par des facteurs environnementaux confondants. Dans son ensemble, cette thèse illustre que, contrairement aux idées reçues, les vecteurs de malaria aviaire ne sont pas toujours permissifs avec leurs parasites. L'environnement apparaît aussi comme un facteur déterminant dans la réponse des vecteurs face à la menace d'infection malarique. Cela pourrait fortement affecter l'épidémiologie de la malaria aviaire.

Variations interindividuelles des performances cognitives et conséquences évolutives chez une population naturelle de mésange charbonnière (Parus major)

Les animaux font face à des changements environnementaux brutaux dus aux modifications de milieux liés à l'activité humaine et aux changements climatiques, et doivent s'ajuster rapidement à leur nouvel environnement. Certains processus cognitifs comme l'innovation et l'apprentissage permettent aux animaux d'intégrer de nouveaux comportements à leur répertoire comportemental (flexibilité comportementale), leur donnant l'opportunité d'intégrer un comportement plus optimal pour s'ajuster. Les performances cognitives varient entre espèces et les individus d'une même population et bien que des études récentes se soient intéressées aux causes des variations interindividuelles des performances cognitives, les conséquences restent peu explorées. Dans cette thèse, les questions des pressions de sélection s'exerçant sur les capacités cognitives sont abordées afin de mieux comprendre l'évolution de ces traits au sein d'une population naturelle de mésange charbonnière Parus major. Un nouveau test de résolution de problème a tout d'abord été présenté à des couples reproducteurs directement en milieu naturel. Les résultats ont montré que les couples les plus performants à résoudre la tâche surpassaient les couples les moins performants sur plusieurs mesures de succès reproducteur. Afin de vérifier que la motivation à nourrir les poussins ne biaisait pas cette relation, la taille de nichée a ensuite été manipulée, ce qui n'a pas affecté la performance subséquente des parents. Les couples innovateurs démontraient un meilleur succès reproducteur quel que soit le changement de la taille de nichée subit, ce qui suggère que cette performance influence bien le succès de reproduction, et non l'inverse. De plus, les couples innovateurs approvisionnaient leurs poussins plus souvent que les couples non innovateurs, suggérant que les innovateurs pourraient exploiter leur habitat de façon plus optimale. Dans un troisième temps, plusieurs caractéristiques morphologiques, dont la coloration des plumes, ont été reliées aux performances de résolution de problème et d'apprentissage. Ces liens, bien que complexes et condition-dépendants, pourraient indiquer un rôle de ces performances lors de la sélection sexuelle. Enfin, afin de tester l'effet du parasite sanguin du paludisme sur les traits comportementaux, un médicament contre le paludisme a été injecté à des femelles reproductrices. Cette injection n'a pas modifié leurs performances cognitives mais a augmenté leur niveau d'activité et d'exploration du nichoir en réponse à la tâche de résolution de problème. Ce parasite sanguin, très présent chez les populations depassereaux, pourrait donc expliquer les variations interindividuelles et interpopulationnelles de certains traits comportementaux en milieu naturel, au même titre que dans nombreux autres systèmes hôte-parasites étudiés. Les travaux présentés dans cette thèse ont permis de détailler pour la première fois la relation entre une performance cognitive et le succès reproducteur chez une population aviaire naturelle, une relation robuste et non influencée par la motivation à nourrir la couvée. Cette performance cognitive est reliée à plusieurs traits morphologiques, mais non à la charge parasitaire. Une meilleure exploitation de l'habitat et habileté à s'occuper des poussins pourrait expliquer cette relation.

Differenzierung von Reviergesängen und mitochondrialem Cytochrom-b in drei ausgewählten Singvogelgattungen (Aves, Passeriformes: Genus Regulus, Genus Seicercus und Parus major)

ZusammenfassungLautäußerungen von Singvögeln (Passeriformes) werden gemeinhin als Träger phylogenetischer Information betrachtet, obwohl direkte Nachweise in vergleichend bioakustischen Studien rar sind. Dieser Thematik widmet sich meine Dissertation am Beispiel dreier Singvogelgruppen: Goldhähnchen (Regulus), Goldbrillenlaubsänger (Seicercus) sowie verwandter Laubsänger (Phylloscopus) und Kohlmeisen (Parus major). Neben der Erhebung bioakustischer Daten wurde für jede Gruppe eine molekulare Phylogenie basierend auf Cytochrom-b-Sequenzen erstellt und für verschiedene akustische Merkmale Homoplasie-indizes berechnet (CI, RI und RC). Die phylogenetisch informativen Gesangsstrukturen innerhalb der Gattungen Regulus und Seicercus/ Phylloscopus sind sämtlich Syntaxmerkmale, zumeist der Gesamtstrophe, seltener von Strophenabschnitten. Bei den Goldhähnchen (Regulus) sind solche Syntaxmerkmale angeboren, Elementmerkmale hingegen sind erlernt und phylogenetisch nicht informativ. Die innerhalb der Kohlmeisen homogene Gesangssyntax ist erst auf höherer taxonomischer Ebene (Gattung Parus) ein informatives Merkmal. Der mittels einer Merkmalsmatrix berechnete akustische Divergenzindex zwischen Taxonpaaren steigt signifikant proportional zur genetischen Distanz. Damit ist erstmalig der Zusammenhang zwischen genetischer und akustischer Differenzierung quantifiziert. Die molekulare Phylogenie erhellt zudem bislang ungeklärte phylogenetische Beziehungen innerhalb aller drei Taxa. Diese werden im Hinblick auf das phylogenetische und das biologische Artkonzept diskutiert. Der Artstatus des Teneriffa-Goldhähnchens (Regulus teneriffae) sowie der bokharensis-Kohlmeisen ist fragwürdig aufgrund ihrer engen Verwandtschaft zu zu einzelnen Subspezies der Wintergoldhähnchen bzw. der Kohlmeisen.

Sperm of colourful males are better protected against oxidative stress

Sperm cells are highly vulnerable to free radicals, and sperm quality and male fertility are critically affected by oxidative stress. Recently, sexual ornaments, particularly carotenoid-based colourful traits, have been proposed to depend on a male's capacity to resist oxidative stress, and thus to signal sperm quality. We conducted an experimental test of this hypothesis on great tits Parus major, in which adults are sexually dichromatic in carotenoid-based breast plumage. We report the first evidence that ornaments and sperm quality may be linked through oxidative stress. When experimentally subjected to oxidative stress resulting from increased workload, less colourful males suffered a greater reduction in sperm motility and swimming ability, and increased levels of sperm lipid peroxidation compared to more colourful males. Moreover, the level of sperm lipid peroxidation was negatively correlated with sperm quality. Finally, carotenoid supplementation increased sperm quality of less colourful males, suggesting that pale males are deficient in carotenoid antioxidants.

Effect of sibling competition and male carotenoid supply on offspring condition and oxidative stress

Early developmental conditions have major implications for an individual's fitness. In species where offspring are born simultaneously, the level of sibling competition for food access is intense. In birds, high sibling competition may subject nestlings to decreased growth rate as a result of limited food and increased levels of oxidative stress through high metabolic activity induced by begging behaviors. We manipulated the level of sibling competition in a natural population of great tits and assessed the consequences for nestling body condition and resistance to oxidative stress. In a full factorial design, we both augmented brood size to increase sibling competition and supplemented the male parents with physiological doses of carotenoids thereby doubling the natural carotenoid intake, aiming at increasing the males' investment in current reproduction and thereby decreasing sibling competition. Nestling body mass was reduced by the brood enlargement and enhanced by the carotenoid supplementation of fathers. Nestling resistance to oxidative stress, measured as total antioxidant defenses in whole blood, was not influenced by the treatments. Because nestlings experience high metabolic activities, an absence of an effect of sibling competition on free radicals production seems unlikely. Nestling body mass decreased and resistance to oxidative stress tended to increase with initial brood size, and hence these correlational effects suggest a trade-off between morphological growth and development of the antioxidant system. However, the result of the experimental treatment did not support this trade-off hypothesis. Alternatively, it suggests that nestling developed compensatory mechanisms that were not detected by our antioxidant capacity measure.

Preparing Offspring for a Dangerous World: Potential Costs of Being Wrong

Adaptive maternal responses to stressful environments before young are born can follow two non-exclusive pathways: either the mother reduces current investment in favor of future investment, or influences offspring growth and development in order to fit offspring phenotype to the stressful environment. Inducing such developmental cues, however, may be risky if the environment changes meanwhile, resulting in maladapted offspring. Here we test the effects of a predator-induced maternal effect in a predator-free postnatal environment. We manipulated perceived predation-risk for breeding female great tits by exposing them to stuffed models of either a predatory bird or a non-predatory control. Offspring were raised either in an environment matching the maternal one by exchanging whole broods within a maternal treatment group, or in a mismatching environment by exchanging broods among the maternal treatments. Offspring growth depended on the matching of the two environments. While for offspring originating from control treated mothers environmental mismatch did not significantly change growth, offspring of mothers under increased perceived predation risk grew faster and larger in matching conditions. Offspring of predator treated mothers fledged about one day later when growing under mismatching conditions. This suggests costs paid by the offspring if mothers predict environmental conditions wrongly.