Jacques Blondel

Feeding ecology and life history variation of the blue tit in Mediterranean deciduous and sclerophyllous habitats.

SummaryHigh variation in laying date and clutch size of the blue tit between a Mediterranean mixed habitat on the mainland, southern France, and a sclerophyllous habitat on the island of Corsica is hypothesized to be related to differences in the food supply. The diet of the nestlings and feeding frequencies were studied using camera nestboxes and electronic chronographs. Food items brought to the nestlings were much more diverse on Corsica than on the mainland, including many fewer caterpillars and a wider range of taxa. However, when expressed as a volume index, prey items were on average larger on Corsica than on the mainland. Feeding frequencies were significantly lower on Corsica. A good correlation was found in both habitats between laying date and the caterpillar peak date, although both the leafing development of oaks and the peak of abundance of caterpillars occurred 3 weeks later in the Corsican sclerophyllous trees than in the mainland deciduous ones. Differences in the feeding ecology of tits between the two habitats are discussed in the light of the evergreen habit, which means that only 30% of leaves are available for phyllophagous insects instead of 100% in deciduous trees. the combination of a late and low food supply in evergreen trees is the best explanation for the differences in breeding traits betwen the two populations.

VARIATION IN BLUE TIT BREEDING PERFORMANCE ACROSS GRADIENTS IN HABITAT RICHNESS

Food supply is widely considered to be a major factor in determining life history traits and reproductive performance of birds. However, large spatial and temporal variation in natural available food supply is not always paralleled by proportional changes in energy demand by breeding birds. This necessarily results in variation in the supply–demand ratio and the amount of food available per unit mass of nestling. Because reproductive performance should respond to increases in available supply as a saturation curve, reaching a plateau above a certain threshold of food supply, we predict that variation in supply should change the intensity of selection on reproductive traits. We first tested this prediction using long-term data on nestling growth and survival in Blue Tits (Parus caeruleus) breeding over a gradient of habitat richness in Corsica, France. This long-term data analysis evaluates the effect of variation in food supply available to breeding tits using three surrogate variables: interannual variation in peak caterpillar abundance (caterpillar frass fall), offset between breeding date and peak caterpillar abundance, and natural variation in clutch size. We also used an experimental brood size manipulation (±3 chicks) to test the effect of varying brood demand on nestling growth. Results of the long-term data analysis show that all three variables affect fledging mass and fledging success in poor habitats, while only fledging mass is affected by variation in offset in rich habitats. Moreover, mean annual fledging success and fledging mass is strongly affected by annual variation in peak caterpillar abundance at low levels of abundance, but these effects disappear when food becomes abundant (saturation threshold level of 373 and 560 mg frass·m−2·d−1 for fledging success and fledging mass, respectively). Brood size manipulations confirm these results. In rich habitats, breeding birds can raise three extra chicks without any apparent effects on chick growth, while in poor habitats, chick growth is significantly reduced with brood enlargement. Our study shows that although food limitation can play an important role, it may not always be the primary force shaping life history traits.

A Thirty-Year Study of Phenotypic and Genetic Variation of Blue Tits in Mediterranean Habitat Mosaics

Abstract In recent years, the study of phenotypic and genetic variation has been enhanced by combining genetic, physiological, demographic, and behavioral components of life histories. Using these new approaches, we address the problem of adaptation to environmental heterogeneity by examining in detail the variation of several fitness-related traits in a small passerine bird, the blue tit, which has been extensively studied in habitat mosaics of the Mediterranean region. The response of blue tits to spatial habitat heterogeneity depends on their range of dispersal relative to the size of habitat patches. Dispersal over short distances leads to local specialization, whereas dispersal over long distances leads to phenotypic plasticity. Gene flow between habitats of different quality may produce local maladaptation and a source–sink population structure. However, when habitat-specific divergent selection regimes are strong enough to oppose the effects of gene flow, local adaptation may arise on a scale that is much smaller than the scale of dispersal.

Molecular phylogeny and the historical biogeography of the warblers of the genus Sylvia (Aves)

A molecular phylogeny based on DNA/DNA hybridization revealed that the Sylvia‐Parisoma complex is monophyletic and includes three main groups of species, the “mid‐European” warblers, the genus Parisoma, and the “eu‐Mediterranean” Sylvia species sensu stricto. The latter can be assigned to three main clusters, a “West‐Mediterranean” group, a “Central‐Mediterranean group”, and an “East‐Mediterranean” group. The radiation of the whole complex is much more ancient than formerly believed. It started ca 12–13 Ma ago and the ancestors of the main extant groups differentiated during the Pliocene. Only speciation events within the “eu‐Mediterranean” lineages occurred during the Pleistocene. The paleoclimatical and paleoecological history of the Mediterranean region is too complicated to provide any evidence for direct relationships between past events and evolutionary steps of these taxa which did not leave any reliable fossil record. However, some major speciation events may be related to well documented climatical crises as well as paleobotanical data. The largely man‐induced extension of matorrals over several millenia presumably extended the range of several species that were formerly much more restricted, which complicates reconstruction of the spatio‐temporal course of speciation.

Nestling diet variation in an insular Mediterranean population of blue tits Parus caeruleus: effects of years, territories and individuals

Mediterranean evergreen forests of Corsica are characterized by relatively high species diversity of arthropods with low population densities. Food is never superabundant for Corsican blue tits Parus caeruleus. This study focused on the composition of the food of blue tit nestlings and especially on two main components, caterpillars and spiders. The nestling diet was studied for two years using 8-mm cameras that automatically took photographs of adult birds with food. The diet was composed of c. 50% caterpillars and c. 30% spiders. There were between-year and between-individual differences in these proportions. In both years of the study the proportion of caterpillars declined during the course of the breeding season. Individual and time effects on prey sizes were also observed. Pairs and individuals were fairly constant in the proportions of prey over the feeding period. Different food items were not brought in runs. These findings suggest that strong food limitation exists on Corsica, which can considerably influence life-history traits of the blue tit.

Adaptive inter-population differences in blue tit life-history traits on Corsica

SummaryFew studies of natural populations have investigated how phenotypic variation across populations relates to key factors in the environment and landscape structure. In the blue tits of southern France, inter-population differences in reproductive life-history traits (e.g. laying date and clutch size) are small, whatever the timing of maximum caterpillar availability, a key factor for offspring survival in tits. These small differences are attributed to gene flow between local populations occupying different habitat types. In contrast, in blue tits on the island of Corsica, we noted large differences in reproductive life-history traits between two populations, where each population is synchronized with the peak-date of caterpillar abundance. These occur over a short geographical distance (25 km). Considering our study within a framework of long-term population studies in tits, our results support the hypothesis that different blue tit populations on Corsica show adaptive differences in life-history traits, and suggest that landscape structure at a small spatial scale can have profound effects on adaptive between-pop illation differentiation in life-history traits that are closely linked with fitness.

Habitat quality as a predictor of spatial variation in blue tit reproductive performance: a multi-plot analysis in a heterogeneous landscape.

Vertebrate studies have rarely investigated the influence of spatial variation in habitat richness on both short-term (breeding) and long-term (offspring recruitment) reproductive performance using simultaneously multi-patch, multi-habitat type and multi-year approaches at landscape level. Here we present results of such an approach using the influence of two oak tree (Quercus ilex, Q. humilis) species on reproductive performance in Corsican blue tits (Parus caeruleus ogliastrae) as a model system. We found that blue tits breeding in rich broad-leaved deciduous patches consistently laid eggs earlier in the season, and produced larger clutches and more fledglings of higher quality, than those breeding in poor evergreen patches. Also, parents, especially males, were in better physical condition in the broad-leaved deciduous than in the evergreen patches. Surprisingly, estimates of long-term effects of reproduction, such as recruitment rates of locally born offspring, did not differ between the two habitat types. Our results suggest that short-term breeding performance and phenotypic quality of both chicks and parents do not necessarily provide reliable information about contributions to following generations at a scale larger than that of the local study plot. Differences in reproductive performance between the two oak habitat types could not be attributed to density-dependent effects, differences in levels of nest predation, or differences in age structure of the birds. We suggest that habitats that are optimal for breeding are not necessarily optimal for survival after the breeding season.

High blowfly parasitic loads affect breeding success in a Mediterranean population of blue tits

Abstract The detrimental effects of ectoparasites on the breeding success of birds have been especially well demonstrated in the case of ectoparasites that affect both chicks and their parents. Since blowfly larvae of the genus Protocalliphora attack only nestling birds, they represent a good model for testing the consequences of parasitism on nestlings. A Corsican population of blue tit suffers extremely high rates of infestation by blowflies, which are suspected to negatively affect young birds. Comparing experimentally deparasitized (treated) and naturally infested (control) broods, we showed that the attack by Protocalliphora causes anaemia and an important disturbance to the chicks. Therefore, we expected that these effects would have a negative impact on body condition and survival in the infested broods. Although we did not find any effect of treatment on fledging success, our predictions were confirmed by lower growth rate, body mass at fledging and tarsus length at fledging in the control compared with the treated group. This suggests that in this population, blowflies decrease the probability of recruitment of young blue tits.

Aromatic plants in nests of the blue tit Cyanistes caeruleus protect chicks from bacteria

Several bird species add fresh fragments of plants which are rich in volatile secondary compounds to their nests. It has been suggested, although never tested, that birds use fresh plants to limit the growth of nest microorganisms. On Corsica, blue tits (Cyanistes caeruleus) incorporate fresh fragments of aromatic plants into their nests. These plants do not reduce infestation by nest ectoparasites, but have been shown to improve growth and condition of chicks at fledging. To understand the mechanisms underlying such benefits, we experimentally tested the effects of these plants on the bacteria living on blue tits. Aromatic plants significantly affected the structure of bacterial communities, in particular reducing bacterial richness on nestlings. In addition, in this population where there is a strong association between bacterial density and infestation by blood-sucking Protocalliphora blow fly larvae, these plants reduced bacterial density on the most infested chicks. Aromatic plants had no significant effect on the bacteria living on adult blue tits. This study provides the first evidence that fresh plants brought to the nests by adult birds limit bacterial richness and density on their chicks.

Local specialization and maladaptation in the Mediterranean blue tit (Parus caeruleus)

Although the evolutionary interplay between gene flow and local adaptation of organisms in heterogeneous environments has been widely discussed from a theoretical point of view, few empirical studies have been designed to test predictions on the consequences of habitat patchiness on the evolution of life history traits. Using blue tits in Mediterranean habitat mosaics as a model, we defined two nested levels of habitat heterogeneity: an inter-regional level which compares two isolated landscapes (mainland, southern France vs the island of Corsica), and an intra-regional level which compares two habitat types within each landscape (deciduous vs evergreen trees). Deciduous habitats are more common than evergreen habitats on the mainland whereas the opposite is true on the island. Results suggest that: (1) on a regional scale, each population is specialized to the more common habitat, i.e. life history traits have evolved in such a way that breeding success is high; (2) in the less common habitats within each landscape, birds are clearly mistimed because they mismatch the best period of food availability, and hence their breeding success is lower; and (3) the density of the populations and the morphometry of the birds support the model of ideal despotic distribution. These results, which are supported by preliminary data on the genetic variation and gene flow of populations in the mainland landscape, are consistent with a source-sink model of population structure within each landscape. They are discussed in the framework of metapopulation theory and habitat selection models.