Sex-specific estimates of dispersal show female philopatry and male dispersal in a promiscuous amphibian, the alpine salamander (Salamandra atra).

Amphibians display wide variations in life-history traits and life cycles that should prove useful to explore the evolution of sex-biased dispersal, but quantitative data on sex-specific dispersal patterns are scarce. Here, we focused on Salamandra atra, an endemic alpine species showing peculiar life-history traits. Strictly terrestrial and viviparous, the species has a promiscuous mating system, and females reproduce only every 3 to 4 years. In the present study, we provide quantitative estimates of asymmetries in male vs. female dispersal using both field-based (mark-recapture) and genetic approaches (detection of sex-biased dispersal and estimates of migration rates based on the contrast in genetic structure across sexes and age classes). Our results revealed a high level of gene flow among populations, which stems exclusively from male dispersal. We hypothesize that philopatric females benefit from being familiar with their natal area for the acquisition and defence of an appropriate shelter, while male dispersal has been secondarily favoured by inbreeding avoidance. Together with other studies on amphibians, our results indicate that a species' mating system alone is a poor predictor of sex-linked differences in dispersal, in particular for promiscuous species. Further studies should focus more directly on the proximate forces that favour or limit dispersal to refine our understanding of the evolution of sex-biased dispersal in animals.

Mating frequency of ant queens with alternative dispersal strategies, as revealed by microsatellite analysis of sperm

In social Hymenoptera (ants, bees, and wasps), the number of males that mate with the same queen affects social and genetic organization of the colony. However, the selective forces leading to single mating in certain conditions and multiple mating in others remain enigmatic. In this study, I investigated whether queens of the wood ant Formica paralugubris adopting different dispersal strategies varied in their mating frequency (the number of males with whom they mated). The frequency of multiple mating was determined by using microsatellite markers to genotype the sperm stored in the spermatheca of queens, and the validity of this method was confirmed by analysing mother-offspring combinations obtained from experimental single-queen colonies. Dispersing queens, which may found new colonies, did not mate with more males than queens that stayed within polygynous colonies, where the presence of numerous reproductive individuals ensured a high level of genetic diversity. Hence, this study provides no support to the hypotheses that multiple mating is beneficial because it increases genetic variability within colonies. Most of the F. paralugubris queens mated with a single male, whatever their dispersal strategy and life history. Moreover, multiple mating had little effect on colony genetic structure: the effective mating frequency was 1.11 when calculated from within-brood relatedness, and 1.13 when calculated from the number of mates detected in the sperm. Hence, occasional multiple mating by F. paralugubris queens may have no adaptive significance.

Ontogeny of Sleeping in Color - Linking melanism to sleep architecture and rhythmicity variation in wild barn owl nestlings

The function of sleep remains unknown. To gain insight into the function of sleep in natural conditions, I assessed variation in sleep architecture and its link with fitness-related phenotypic traits. I considered melanin-based coloration because its underlying genetic basis is very well known giving an opportunity to examine whether some genes pleiotropically regulate both coloration and sleep. The melanocortin system is known to generate covariation between melanin-based coloration and other phenotypes like behaviour, physiology and life history traits. I investigated whether this system of genes could participate in the co-expression of coloration and sleep. I carried out a study with nestling barn owls (Tyto alba) in order to tackle the potential link between variation in color traits and the ontogeny of sleep under natural conditions. For this I established a suitable method for recording the brain activity (electroencephalogram) of owls in nature. Birds are especially interesting, because they convergently evolved sleep states similar to those exhibited by mammals. As in mammals, I found that in owlets time spent in rapid eye movement (REM) sleep declines with age, a relationship thought to eflect developmental changes in the brain. Thus this developmental trajectory appears to reflect a fundamental feature of sleep. Additionally, I discovered an association between a gene involved in melanism expressed in the feather follicles (proprotein convertase subtilisin/kexin type 2, PCSK2) and the age-related changes in sleep in the brain. Nestlings with higher expression levels of PCSK2 showed a more precocial pattern of sleep development and a higher degree of melanin-based coloration compared to nestlings with lower PCSK2 expression. Also sleep architecture and the development of rhythmicity in brain and physical activity was related to plumage traits of the nestlings and their biological parents. This pattern during ontogeny might reflect differences in life l history strategies, antipredator behaviour and developmental pace. Therefore, differently colored individuals may differentially deal with trade-offs between the costs and benefits of sleep which in turn lead to differences in brain organization and ultimately fitness. These results should stimulate evolutionary biologists to consider sleep as a major life history trait. Résumé La fonction du sommeil reste inconnue. Afin d'acquérir une meilleur compréhension de la fonction du sommeil dans les conditions naturelles, j'ai analysé la variation dans l'architecture du sommeil et son lien avec d'autres traits phénotypiques liés au succès reproducteur (fitness). J'ai choisi et examiné la coloration mélanique, car ses bases génétiques sont bien connues et il est ainsi possible d'étudier si certains gènes, de façon pléiotropique régulent à la fois la coloration et le sommeil. J'ai exploré si ce système génétique était impliqué dans la co-expression de la coloration et du sommeil. J'ai effectué mon étude sur des poussins de chouette effraie (Tyto alba) en condition naturelle, pour rechercher ce lien potentiel entre la variation de la coloration et l'ontogenèse du sommeil. Dans ce but, j'ai établi une méthodologie permettant d'enregistrer l'activité cérébrale (électroencéphalogramme) des chouettes dans la nature. Les oiseaux sont particulièrement intéressants car ils ont développé, par évolution convergente, des phases de sommeil similaires à celles des mammifères. De manière semblable à ce qui a été montré chez les mammifères, j'ai découvert que le temps passé dans le sommeil paradoxal diminue avec l'âge des poussins. On pense que ceci est dû aux changements développementaux au niveau du cerveau. Cette trajectoire développementale semble refléter une caractéristique fondamentale du sommeil. J'ai également découvert une association entre l'un des gènes impliqué dans le mélanisme, exprimé dans les follicules plumeux (proprotein convertase subtilisin/kexin type 2, PCSK2), et les changements dans la structure du sommeil avec l'âge. Les poussins ayant un niveau d'expression génétique élevé de la PCSK2 présentent une structure du sommeil plus précoce et un taux de coloration dû à la mélanine plus élevé que des poussins avec un niveau d'expression moindre de la PCSK2. L'architecture du sommeil et le développement de la rythmicité dans le cerveau ainsi que l'activité physique sont également liés à la coloration des plumes des poussins et pourraient ainsi refléter des différences de stratégies d'histoire de vie, de comportements anti-prédateur et de vitesses développementales. Ainsi, des individus de coloration différente sembleraient traiter différemment les coûts et les bénéfices du sommeil, ce qui aurait des conséquences sur l'organisation cérébrale et pour finir, sur le succès reproducteur. Ces résultats devraient encourager les biologistes évolutionnistes à considérer le sommeil comme un important trait d'histoire de vie. Zusammenfassung Die Funktion von Schlaf ist noch unbekannt. Um mehr Einsicht in diese unter natürlichen Bedingungen zu bekommen, habe ich die Variation in der Schlafarchitektur und die Verknüpfung mit phänotypischen Merkmalen, die mit der Fitness zusammenhängen, studiert. Ich habe mir melanin-basierte Färbung angesehen, da die zugrunde liegende genetische Basis bekannt ist und somit die Möglichkeit gegeben ist, zu untersuchen, ob einige Gene beides regulieren, Färbung und Schlaf. Das melanocortin System generiert eine Kovariation zwischen melanin-basierter Färbung und anderen phänotypischer Merkmale wie Verhalten, Physiologie und Überlebensstrategien. Ich habe untersucht, ob dieses Gensystem an einer gleichzeitigen Steuerung von Färbung und Schlaf beteiligt ist. Dazu habe ich Schleiereulen (Tyto alba) studiert um einen möglichen Zusammenhang zwischen der Variation in der Pigmentierung und der Entwicklung des Schlafs unter natürlichen Bedingungen zu entdecken. Für diese Studie entwickelte ich eine Methode um die Gehirnaktivität (Elektroenzephalogramm) bei Eulen in der Natur aufzunehmen. Vögel sind besonders interessant, da sie die gleichen Schlafstadien aufweisen wie Säugetiere und diese unabhängig konvergent entwickelt haben. Genauso wie bei Säugetieren nahm die Dauer des sogenannten ,,rapid eye movement" (REM) - Schlafes mit zunehmendem Alter ab. Es wird angenommen, dass dieser Zusammenhang die Entwicklung des Gehirns widerspiegelt. Daher scheint dieses Entwicklungsmuster ein fundamentaler Aspekt von Schlaf zu sein. Zusätzlich entdeckte ich einen Zusammenhang zwischen der Aktivität eines Gens in den Federfollikeln (proprotein convertase subtilisin/kexin type 2, PCSK2), das für die Ausprägung schwarzer Punkte auf den Federn der Eulen verantwortlich ist, und den altersabhängigen Änderungen im Schlafmuster im Gehirn. Küken mit höherer Aktivität von PCSK2 zeigten eine frühreifere Schlafentwicklung und eine dunklere Färbung als Küken mit niedriger PCSK2 Aktivität. Die Architekture des Schlafes und die Entwicklung der Rhythmik im Gehirn und die der physischen Aktivität ist mit der Färbung des Gefieders von den Küken und ihren Eltern verknüpft. Dieses Muster während der Entwicklung kann Unterschiede in Überlebensstrategien, Feindabwehrverhalten und in der Entwicklungsgeschwindigkeit reflektieren. Unterschiedlich gefärbte Individuen könnten unterschiedliche Strategien haben um zwischen den Kosten und Nutzen von Schlaf zu entscheiden, was zu Unterschieden in der Gehirnstruktur führen kann und letztendlich zur Fitness. Diese Ergebnisse sollten Evolutionsbiologen stimulieren Schlaf als einen wichtigen Bestandteil des Lebens zu behandeln.

The effect of innovation and sex-specific migration on neutral cultural differentiation

Studies of behaviour are increasingly focusing on acquisition of traits through cultural inheritance. Comparison of patterns of spatial population structure (FST) between neutral genetic loci and behavioural or cultural traits can been used to test hypotheses about demography, life history, and the mechanisms of inheritance/transmission of these traits in humans, chimpanzees and other animals. Here, we develop analytical expectations to show how FST in cultural traits can differ strongly from that measured at neutral genetic markers if migration is largely restricted to one sex but social learning is predominantly modelled on the other (e.g. males migrate, females serve as models for cultural traits), if one individual is the learning model for many, or if rates of innovation (individual learning) are high or rates of social learning are low. We discuss how comparisons of FST between genetic loci and behavioural traits can be applied to evaluate the importance of innovation in shaping patterns of cultural differentiation, as even low rates of innovation can considerably reduce FST, relative to observed structure at neutral genetic loci. Our results also suggest that differentiation in neutral cultural traits should occur over much smaller scales in species with male migration and female enculturation (or the reverse).

Anthropogenic disturbance as a driver of microspatial and microhabitat segregation of cytotypes of Centaurea stoebe and cytotype interactions in secondary contact zones.

BACKGROUND AND AIMS: In a mixed-ploidy population, strong frequency-dependent mating will lead to the elimination of the less common cytotype, unless prezygotic barriers enhance assortative mating. However, such barriers favouring cytotype coexistence have only rarely been explored. Here, an assessment is made of the mechanisms involved in formation of mixed-ploidy populations and coexistence of diploid plants and their closely related allotetraploid derivates from the Centaurea stoebe complex (Asteraceae). METHODS: An investigation was made of microspatial and microhabitat distribution, life-history and fitness traits, flowering phenology, genetic relatedness of cytotypes and intercytotype gene flow (cpDNA and microsatellites) in six mixed-ploidy populations in Central Europe. KEY RESULTS: Diploids and tetraploids were genetically differentiated, thus corroborating the secondary origin of contact zones. The cytotypes were spatially segregated at all sites studied, with tetraploids colonizing preferentially drier and open microhabitats created by human-induced disturbances. Conversely, they were rare in more natural microsites and microsites with denser vegetation despite their superior persistence ability (polycarpic life cycle). The seed set of tetraploid plants was strongly influenced by their frequency in mixed-ploidy populations. Triploid hybrids originated from bidirectional hybridizations were extremely rare and almost completely sterile, indicating a strong postzygotic barrier between cytotypes. CONCLUSIONS: The findings suggest that tetraploids are later immigrants into already established diploid populations and that anthropogenic activities creating open niches favouring propagule introductions were the major factor shaping the non-random distribution and habitat segregation of cytotypes at fine spatial scale. Establishment and spread of tetraploids was further facilitated by their superior persistence through the perennial life cycle. The results highlight the importance of non-adaptive spatio-temporal processes in explaining microhabitat and microspatial segregation of cytotypes.

Fecundity and survival in relation to resistance to oxidative stress in a free-living bird.

Major life history traits, such as fecundity and survival, have been consistently demonstrated to covary positively in nature, some individuals having more resources than others to allocate to all aspects of their life history. Yet, little is known about which resources (or state variables) may account for such covariation. Reactive oxygen species (ROS) are natural by-products of metabolism and, when ROS production exceeds antioxidant defenses, organisms are exposed to oxidative stress that can have deleterious effects on their fecundity and survival. Using a wild, long-lived bird, the Alpine Swift (Apus melba), we examined whether individual red cell resistance to oxidative stress covaried with fecundity and survival. We found that males that survived to the next breeding season tended to be more resistant to oxidative stress, and females with higher resistance to oxidative stress laid larger clutches. Furthermore, the eggs of females with low resistance to oxidative stress were less likely to hatch than those of females with high resistance to oxidative stress. By swapping entire clutches at clutch completion, we then demonstrated that hatching failure was related to the production of low-quality eggs by females with low resistance to oxidative stress, rather than to inadequate parental care during incubation. Although male and female resistance to oxidative stress covaried with age, the relationships among oxidative stress, survival, and fecundity occurred independently of chronological age. Overall, our study suggests that oxidative stress may play a significant role in shaping fecundity and survival in the wild. It further suggests that the nature of the covariation between resistance to oxidative stress and life history traits is sex specific, high resistance to oxidative stress covarying primarily with fecundity in females and with survival in males.

The adaptive function of melanin-based coloration in the tawny owl (Strix aluco) ; the proximate role of the melanocortin system

Colour polymorphism is common in wild population. One of the main questioning of evolutionary biologists is to understand how different colour variants could have evolved and be maintained in fluctuating environments, a selective process that forces individuals to constantly adapt their strategies in order to survive. This issue is particularly true for traits that are genetically inherited. Natural selection erodes genotypes with lowest fitness (less adapted), reducing in turn global genetic variation within population. In this context, the study of the evolution and maintenance of melanin- based coloration is relevant since inter-individual variation in the deposition of these pigments is common in animal and plant kingdoms and is under strong genetic control. In this thesis, I focus on the specific case of the tawny owl (Strix aluco), a species displaying continuous variation in reddish pheomelanin-based coloration. Interestingly, empirical studies highlighted covariations between melanin-based coloration and important behavioural, physiological and life history traits. Recently, a genetic model pointed out the melanocortin system and their pleiotropic effects as a potential regulator of these covariations. Accordingly, this PhD thesis further investigates colour-specific behavioural, physiological, or life history strategies, while examining the proximate mechanisms underlying these reaction norms. We found that differently coloured tawny owls differently resolve fundamental trade-off between offspring number and quality (Chapter 1), light melanic individuals producing many low- quality offspring and dark, melanic ones producing few high-quality offspring. These reproductive strategies are likely to induce alternative physiological constraints. Indeed, we demonstrated that light melanic individuals produced higher levels of reactive oxygen species (ROS, Chapter 2), but also expressed higher levels of antioxidant (GSH, Chapters 2 & 3). Interestingly, we showed that light melanic breeding females could modulate their POMC prohormone levels according to the environmental conditions, while dark reddish ones produced constant levels of this prohormone {Chapter 4). Finally, we highlighted colour-specific patterns of prohormone convertase 1 (PCI) gene expression (Chapter 5), an enzyme responsible for POMC prohormone processing to ACTH and a- MSH, for instance. Altogether, these results provide strong evidence of colour-specific strategies, light and melanic tawny owls better coping with stressful and relaxed environments, respectively. Variation in melanin-based coloration is likely to be maintained by the heterogeneity of our study area and strong environmental stochasticity within and between years, these process favouring differently coloured tawny owls at different periods of time. From a proximate point of view, this PhD thesis supports the hypothesis that covariations between phenotypic traits and melanin-based coloration stems from the melanocortin system, especially the fundamental role of POMC gene expression and its processing to melanocortin peptides. - Le polymorphisme de couleur est une variation phénotypique très fréquente dans la nature. En biologie évolutive, une des problématiques clés est donc de comprendre comment différent morphes de couleur peuvent être apparus et maintenus au cours du temps dans des environnements aussi variables que les nôtres, surtout que ces fluctuations forcent ces morphes à s'adapter constamment pour assurer leur survie. Cette thématique est particulièrement réelle lorsque les variations phénotypiques sont héréditaires et donc sous forte influence génétique. La sélection naturelle a en effet le pouvoir d'éroder rapidement la variation génétique en éliminant les génotypes mal adaptés. Dans ce sens, l'étude de l'évolution, et de la maintenance de la coloration mélanique est donc tout à fait pertinente car la variation de coloration entre individus est très répandue à travers les règnes animal et végétal et sous forte influence génétique. Dans cette thèse, je me suis concentré sur le cas spécifique de la chouette hulotte (Strix aluco), une espèce présentant une variation continue dans la déposition de pigments pheomélaniques roux. De précédentes études ont déjà montré que cette variation de coloration était associée avec des variations de traits comportementaux, physiologiques ou d'histoire de vie. Récemment, une étude a souligné l'importance du système des mélanocortines et de leurs effets pléiotropes dans la régulation de ces covariations. En conséquence, cette thèse de doctoral a pour but d'étudier un peu plus les stratégies comportementales, physiologiques ou d'histoire de vie spécifiques à chaque morphe de couleur, tout en examinant un peu plus les mécanismes proximaux potentiellement à la base de ces normes de réactions. Nous constatons tout d'abord que les morphes de couleurs étaient associés à différentes stratégies dans la résolution de compromis telle que la production de beaucoup de jeunes ou des jeunes de qualité (Chapitre 1). Les morphes gris (dit peu mélaniques) ont tendance à produire beaucoup de jeunes mains de moindre qualité, alors que les morphes roux (dit fortement mélaniques) produisent moins de jeunes mais de meilleure qualité. Ces stratégies sont susceptibles alors d'induire certaines contraintes physiologiques. Par exemple, nous montrons que les morphes gris produisent plus de dérivés réactifs de l'oxygène (ROS, Chapitre 2), mais aussi plus d'antioxydants (GSH, Chapitres 2 & 3). Nous montrons ensuite que les femelles grises ont une plus grande capacité à moduler leur niveau de POMC prohormone dans le sang en fonction des conditions environnementales, alors que les femelles rousses gardent un niveau constant (Chapitre 4). Finalement, nous démontrons que les patterns d'expression du gène codant pour la prohormone convertase 1 varient chez des jeunes issus de parents gris ou roux (Chapitre 5). Ceci est particulièrement intéressant car cette enzyme permet de scinder la POMC prohormone en plusieurs peptides importants tels que l'ACTH ou l'a-MSH. En conclusion, ces résultats démontrent qu'il y a bel et bien des stratégies évolutives différentes entre les morphes de couleurs, les chouettes hulottes grises et rousses étant respectivement plus adaptés à des environnements stressants ou favorables. L'hétérogénéité de notre zone d'étude et la stochasticité environnementale qui caractérise ses habitats pourraient donc agir comme une source de sélection temporelle, laquelle favoriserait les différents morphes de couleurs à diverses périodes. D'un point de vue plus proximale maintenant, cette thèse de doctorat soutient l'hypothèse que les covariations observées entre la coloration mélanique et des traits phénotypiques importants sont modulées par les effets pléiotropes du système des mélanocortines, et met en avant le rôle prépondérant que pourrait jouer l'expression du gène POMC et sa post traduction en mélanocortines.

MHC class I expression dependent on bacterial infection and parental factors in whitefish embryos (Salmonidae).

Ecological conditions can influence not only the expression of a phenotype, but also the heritability of a trait. As such, heritable variation for a trait needs to be studied across environments. We have investigated how pathogen challenge affects the expression of MHC genes in embryos of the lake whitefish Coregonus palaea. In order to experimentally separate paternal (i.e. genetic) from maternal and environmental effects, and determine whether and how stress affects the heritable variation for MHC expression, embryos were produced in full-factorial in vitro fertilizations, reared singly, and exposed at 208 degree days (late-eyed stage) to either one of two strains of Pseudomonas fluorescens that differ in their virulence characteristics (one increased mortality, while both delayed hatching time). Gene expression was assessed 48 h postinoculation, and virulence effects of the bacterial infection were monitored until hatching. We found no evidence of MHC class II expression at this stage of development. MHC class I expression was markedly down-regulated in reaction to both pseudomonads. While MHC expression could not be linked to embryo survival, the less the gene was expressed, the earlier the embryos hatched within each treatment group, possibly due to trade-offs between immune function and developmental rate or further factors that affect both hatching timing and MHC expression. We found significant additive genetic variance for MHC class I expression in some treatments. That is, changes in pathogen pressures could induce rapid evolution in MHC class I expression. However, we found no additive genetic variance in reaction norms in our study population.

Physiologie des sexués femelles de fourmis (Hymenoptera; Formicidae) en relation avec le mode de fondation

A study was undertaken to determine if there was a relation between the mode of colony founding in ants and the physiology of the new queens produced, in which mature gynes of 24 ant species were examined. Gynes of species utilizing independent colony founding had a far higher relative fat content (X±SD; 54±6%)(g fat/g dry weight) than gynes of species employing dependent colony founding(19±8%). Dimorphism between queens and workers was significantly higher in species employing independent colony founding. Thus independent colony founding not only results in production of queens with a relatively higher fat content and therefore with a higher energy content per g, but also results in the production of larger queens (in comparison with worker size). Of species employing independent colony founding, 80% were monogynous, whereas only 11% of the species employing dependent colony founding were monogynous. These results are discussed with regard to the social structure and life-history of ant species.

Variation in thermal performance and reaction norms among populations of Drosophila melanogaster.

The major goal of evolutionary thermal biology is to understand how variation in temperature shapes phenotypic evolution. Comparing thermal reaction norms among populations from different thermal environments allows us to gain insights into the evolutionary mechanisms underlying thermal adaptation. Here, we have examined thermal adaptation in six wild populations of the fruit fly (Drosophila melanogaster) from markedly different natural environments by analyzing thermal reaction norms for fecundity, thorax length, wing area, and ovariole number under ecologically realistic fluctuating temperature regimes in the laboratory. Contrary to expectation, we found only minor differences in the thermal optima for fecundity among populations. Differentiation among populations was mainly due to differences in absolute (and partly also relative) thermal fecundity performance. Despite significant variation among populations in the absolute values of morphological traits, we observed only minor differentiation in their reaction norms. Overall, the thermal reaction norms for all traits examined were remarkably similar among different populations. Our results therefore suggest that thermal adaptation in D. melanogaster predominantly involves evolutionary changes in absolute trait values rather than in aspects of thermal reaction norms.

Natural malaria infection reduces starvation resistance of nutritionally stressed mosquitoes.

In disease ecology, there is growing evidence that environmental quality interacts with parasite and host to determine host susceptibility to an infection. Most studies of malaria parasites have focused on the infection costs incurred by the hosts, and few have investigated the costs on mosquito vectors. The interplay between the environment, the vector and the parasite has therefore mostly been ignored and often relied on unnatural or allopatric Plasmodium/vector associations. Here, we investigated the effects of natural avian malaria infection on both fecundity and survival of field-caught female Culex pipiens mosquitoes, individually maintained in laboratory conditions. We manipulated environmental quality by providing mosquitoes with different concentrations of glucose-feeding solution prior to submitting them to a starvation challenge. We used molecular-based methods to assess mosquitoes' infection status. We found that mosquitoes infected with Plasmodium had lower starvation resistance than uninfected ones only under low nutritional conditions. The effect of nutritional stress varied with time, with the difference of starvation resistance between optimally and suboptimally fed mosquitoes increasing from spring to summer, as shown by a significant interaction between diet treatment and months of capture. Infected and uninfected mosquitoes had similar clutch size, indicating no effect of infection on fecundity. Overall, this study suggests that avian malaria vectors may suffer Plasmodium infection costs in their natural habitat, under certain environmental conditions. This may have major implications for disease transmission in the wild.

Additive effects of ectoparasites over reproductive attempts in the long-lived Alpine swift

1. Parasitism is a non-negligible cost of reproduction in wild organisms, and hosts are selected to partition resources optimally between current and future reproduction. While parents can compensate for the cost of parasitism by increasing their current reproductive investment, such change in resource allocation is expected to carry-over costs on future reproduction. 2. Life history theory predicts that because long-lived organisms have a high residual reproductive value, they should be more reluctant to increase parental effort in response to parasites. Also, when rearing successive infested broods, the cost of parasitism can cumulate over the years and hence be exacerbated by past infestations. 3. We tested these two predictions in the alpine swift Apus melba, a long-lived colonial bird that is infested intensely by the nest-based blood sucking louse-fly Crataerina melbae. For this purpose, we manipulated ectoparasite load over 3 consecutive years and measured reproductive parameters in successive breeding attempts of adults assigned randomly to 'parasitized' and 'deparasitized' treatments. 4. In current reproduction, fathers of experimentally parasitized broods produced a similar number of offspring as fathers from the deparasitized treatment, but the rearing period was prolonged by 4 days. Fathers that were assigned to the parasitized treatment in year x produced significantly fewer fledglings the following year x + 1 than those of the deparasitized treatment. The number of young produced by fathers in year x + 1 was correlated negatively with the number of days they cared for their brood in the previous year x. We also found a significant interaction between treatments performed over 2 successive years, with fathers of parasitized broods suffering a larger fitness loss if in the past they had already cared for a parasitized brood rather than for a deparasitized one. Similar effects of parasitism, although partly non-significant (0.05 < P-values > 0.10), were found in mothers. 5. Altogether, our results show that parasites can modify resource allocation between current and future reproduction in long-lived hosts, and that the cost of parasitism can cumulate over the years. It emphasizes the fact that effects of parasites can depend on past infestations and become apparent in future reproduction only.

Group selection and kin selection: two concepts but one process.

In a recent paper, Traulsen and Nowak use a multilevel selection model to show that cooperation can be favored by group selection in finite populations [Traulsen A, Nowak M (2006) Proc Natl Acad Sci USA 103:10952-10955]. The authors challenge the view that kin selection may be an appropriate interpretation of their results and state that group selection is a distinctive process "that permeates evolutionary processes from the emergence of the first cells to eusociality and the economics of nations." In this paper, we start by addressing Traulsen and Nowak's challenge and demonstrate that all their results can be obtained by an application of kin selection theory. We then extend Traulsen and Nowak's model to life history conditions that have been previously studied. This allows us to highlight the differences and similarities between Traulsen and Nowak's model and typical kin selection models and also to broaden the scope of their results. Our retrospective analyses of Traulsen and Nowak's model illustrate that it is possible to convert group selection models to kin selection models without disturbing the mathematics describing the net effect of selection on cooperation.

Natural selection on fecundity variance in subdivided populations: kin selection meets bet hedging.

In a series of seminal articles in 1974, 1975, and 1977, J. H. Gillespie challenged the notion that the "fittest" individuals are those that produce on average the highest number of offspring. He showed that in small populations, the variance in fecundity can determine fitness as much as mean fecundity. One likely reason why Gillespie's concept of within-generation bet hedging has been largely ignored is the general consensus that natural populations are of large size. As a consequence, essentially no work has investigated the role of the fecundity variance on the evolutionary stable state of life-history strategies. While typically large, natural populations also tend to be subdivided in local demes connected by migration. Here, we integrate Gillespie's measure of selection for within-generation bet hedging into the inclusive fitness and game theoretic measure of selection for structured populations. The resulting framework demonstrates that selection against high variance in offspring number is a potent force in large, but structured populations. More generally, the results highlight that variance in offspring number will directly affect various life-history strategies, especially those involving kin interaction. The selective pressures on three key traits are directly investigated here, namely within-generation bet hedging, helping behaviors, and the evolutionary stable dispersal rate. The evolutionary dynamics of all three traits are markedly affected by variance in offspring number, although to a different extent and under different demographic conditions.

Why is the cladoceran Simocephalus vetulus (Müller) not a 'bang-bang strategist '? A critique of the optimal-body-size model

Lynch's (1980a) optimal-body-size model is designed to explain some major trends in cladoceran life histories; in particular the fact that large and littoral species seem to be bang-bang strategists (they grow first and the reproduce) whereas smaller planktonic species seem to be intermediate strategists (they grow and reproduce simultaneously). Predation is assumed to be an important selective pressure for these trends. Simocephalus vetulus (Müller) does not fit this pattern; being a littoral and relatively large species but an intermediate strategist. As shown by computer simulations, this species would reduce its per capita rate of increase by adopting the strategy predicted by the optimal-body-size model. Two aspects of the model are criticized: (1) the optimization criterion is shown to be incorrect and (2) the prediction of an intermediate strategy is not justified. Structural constraints are suggested to be responsible for the intermediate strategy of S.vetulus. Biotic interactions seem to have little effect on the observed life-history patterns of this species.