Sexual size dimorphism in the great tit (Parus major) in relation to history and current selection

We investigated the possible causes of the evolution of sexual size and shape dimorphism in the great tit (Parus major) by using two different approaches. First, we used the equilibrium approach, i.e. analysing current selection to see whether it was possible to find directional selection in the direction of the dimorphism, or stabilising selection maintaining dimorphism at its current level. Second, we used the historical approach, i.e. putting the degree of dimorphism in a phylogenetic perspective to analyse what kind of changes (if any) have occurred. This was carried out in the following way: (i) we described the level of sexual dimorphism in a population of Swedish great tits by means of path model. (ii) We used the path model design to analyse survival and reproductive selection in this population. (iii) We compared the level of dimorphism in relation to size in the great tit with that of the closest congener, the blue tit P. caeruleus. (iv) We compared the amount of interspecific morphological variation with that which would be expected under a drift model. We found no evidence of either stabilising or directional survival or reproductive selection. Size and shape variation in the great tit seemed unrelated to fitness in adults. Dimorphism was somewhat greater in the great tit compared to the blue tit, but only with an amount predictable by its larger size. In terms of phenotypic standard deviations, the great tit was not more dimorphic than the blue tit, although it was larger. The amount of interspecific variance with regard to size was lower or equal to that expected by the drift model, showing that long-term directional selection for an increase in size and dimorphism is improbable. These results agree with recent theoretical findings that size and dimorphism should be related and that strong conservatism with regard to dimorphism is to be expected. They also agree with the view that in equilibrium populations, fitness components (if there are many of them) should appear neutral with regard to total fitness.     

Asymmetric competition between two tit species: a reciprocal removal experiment

1. In a 5-year field experiment, competition for food was tested between great tit ( Parus major L.) and blue tit ( P. caeruleus L.), two common hole-nesters in Central Europe. Experimental ('allopatric') populations of both species were created during the breeding seasons by preventing the nesting or egg laying of one of the competing species.
2. An asymmetric relationship was found between the two tits; blue tits were more successful in the competitive interaction. Detectable effects were found only in nestling condition. Great tits raised lighter nestlings when breeding sympatrically with blue tits.
3. A possible mechanism is suggested that could be responsible for the different competitive abilities of the two species; blue tits are more effective in utilizing the most abundant size categories of caterpillar food supply than great tits.     

Similarity of growth among Great tits (Parus major) and Blue tits (P. caeruleus)

Differences in morphology among species are proximately caused by changes in the ontogeny of individuals. It is therefore of importance to analyse possible differences in growth parameters among closely related species in order to understand what parameters are most and least likely, respectively, to change in evolution. In this paper I analyse growth in two closely related sympatric species, namely Great tit, Parus major, and Blue tit, P. caeruleus. The former is considerably larger than the latter in all external traits. The growth rates of the two species were found to be very similar for all traits, thus excluding differences in growth rate as a potential cause of evolutionary size changes. Offset of growth occurred at relatively similar times in the two species, excluding this factor as a major cause of the final size differences. However, size differences at hatching were pronounced and remained so throughout ontogeny, pointing to initial size (egg size or hatching size) as the target of factors promoting change. Bivariate allometric relations of traits vs. body size (mass) were similar between the two species at all ontogenetic stages. There was a high correlation among traits especially at intermediate age stages (5 and 8 days), but these correlations became weaker at older age and approached the low pattern of integration found in adults. All this suggests the operation of a general growth factor affecting all parts of the phenotype simultaneously, which has its major influence at the time of maximal growth. If closely related species in general have highly similar growth patterns, strong evolutionary allometry as found in many avian taxa is to be expected.     

The extended Moran effect and large-scale synchronous fluctuations in the size of great tit and blue tit populations

1. Synchronous fluctuations of geographically separated populations are in general explained by the Moran effect, i.e. a common influence on the local population dynamics of environmental variables that are correlated in space. Empirical support for such a Moran effect has been difficult to provide, mainly due to problems separating out effects of local population dynamics, demographic stochasticity and dispersal that also influence the spatial scaling of population processes. Here we generalize the Moran effect by decomposing the spatial autocorrelation function for fluctuations in the size of great tit Parus major and blue tit Cyanistes caeruleus populations into components due to spatial correlations in the environmental noise, local differences in the strength of density regulation and the effects of demographic stochasticity. 2. Differences between localities in the strength of density dependence and nonlinearity in the density regulation had a small effect on population synchrony, whereas demographic stochasticity reduced the effects of the spatial correlation in environmental noise on the spatial correlations in population size by 21.7% and 23.3% in the great tit and blue tit, respectively. 3. Different environmental variables, such as beech mast and climate, induce a common environmental forcing on the dynamics of central European great and blue tit populations. This generates synchronous fluctuations in the size of populations located several hundred kilometres apart. 4. Although these environmental variables were autocorrelated over large areas, their contribution to the spatial synchrony in the population fluctuations differed, dependent on the spatial scaling of their effects on the local population dynamics. We also demonstrate that this effect can lead to the paradoxical result that a common environmental variable can induce spatial desynchronization of the population fluctuations. 5. This demonstrates that a proper understanding of the ecological consequences of environmental changes, especially those that occur simultaneously over large areas, will require information about the spatial scaling of their effects on local population dynamics.     

山地次生林冬季鸟类集群的初步研究

通过山地次生林两冬季鸟类集群的研究表明,集群的鸟类主要是由大山雀、沼泽山雀和普通Ｓｈｉ组成;每群个体数为７．４１－１０．６３只;据群的行为（单种群、多种集群和单独个体）分析,大多数种类是单独个体活动,冬季鸟类集群大小和组成有空间的变化。气候因子影响群的形成和行为。     

Isolation and characterization of 12 tetranucleotide repeat microsatellite loci from the green‐backed tit (Parus monticolus)

From a partial genomic library enriched for GATA short tandem repeats, we developed 12 polymorphic microsatellite loci from the green‐backed tit (Parus monticolus). We characterized these loci by genotyping 30 adult individuals with unknown relationship. The number of alleles ranged from four to 17 per locus (mean = 9.3 alleles) and the observed heterozygosity for each locus ranged from 0.633 to 0.933 (mean = 0.789). All loci conformed to Hardy–Weinberg expectations. Four of 66 possible pairwise comparisons between loci showed significant gametic disequilibrium.     

The diet of great tit nestlings: Comparing observation records and stable isotope analyses

The diet of wild animals has been studied using many different strategies, approaches and methods in recent decades. In this regard, stable isotopes analysis (SIA) is becoming a widespread tool, but no study has yet, to our knowledge, compared diet estimations from SIA with direct observations of the diet of passerine nestlings. Accordingly, our aim was to test the predictive power of SIA for this purpose and identify potential confounding factors such as habitat effects. To do this, we compared isotopic signatures of δ¹³C and δ¹⁵N in the feathers of great tit (Parus major) nestlings, and the corresponding estimates of their diet based on stable isotope mixing models, with prey proportions delivered by their parents obtained through video-recordings. Between-nest differences in isotopic signatures of δ¹⁵N were larger than within-nest differences. We found that δ¹⁵N signatures of nestling feathers correlated positively with the proportion of spiders and negatively with the proportion of caterpillars in the nestlings’ diet, the most important prey types. On the other hand, between-nest and within-nest differences in δ¹³C ratios were of similar magnitude and δ¹³C ratios correlated mainly with the proportion of trees surrounding nest-boxes that were Quercus spp. Estimates of diet composition based on mixing models correlated with the observed nestling diet, yet effect sizes were quite low. Although mixing models are commonly used to ascertain diets, our data show that they can provide valuable information on the relative intake of prey types from different trophic levels; but when complex dietary patterns are recorded (e.g. due to the confounding effects of habitat and/or temporal variation) it can be difficult to draw firm conclusions about diet composition.     

Keypeck conditioning with reinforcements in two different locations in thrush,tit and sparrow species

Twenty-one individuals each of the species great tits $\text{(Parus major),}$ blue tits $\text{(Parus caeruleus),}$ marsh tits $\text{(Parus palustris),}$ blackbirds $\text{(Turdus merula),}$ songthrushes $\text{(Turdus philomelos),}$ tree sparrows $\text{(Passer montanus)}$ and house sparrows $\text{(Passer domesticus)}$ were conditioned to a keypeck response by means of food reinforcement. The site where the reward was given was then changed. The number of reinforcements were then counted until the keypeck response followed immediately (within 10 seconds) on three successive occasions, and a comparison was made between species. Of the species studied the blackbird needed significantly fewer instances of reinforcement in order to re-establish an immediate keypeck response. A significant difference was also seen between the performance of the great tit and marsh tit, as well as between the blue tit and the marsh tit. No significant difference was found between the performance of the house sparrow and the tree sparrow.     

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

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. Received: 6 May 1997 / Accepted: 14 July 1997     

Isolation and characterization of 16 microsatellite loci in the Great Tit Parus major

Six dinucleotide, three trinucleotide and seven tetranucleotide microsatellite loci developed for the great tit Parus major are presented. Thirty individual birds were screened at each locus. Loci were polymorphic (four to 19 alleles per locus). These markers provide a system to study paternity, genetic diversity in natural populations, gene flow, dispersal and inbreeding.     

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.     

Isolation and characterization of 23 microsatellite loci in the hen flea Ceratophyllus gallinae

Nineteen dinucleotide, two trinucleotide and two tetranucleotide microsatellite loci developed for the hen flea Ceratophyllus gallinae are presented. Twenty fleas were screened at each locus. Loci were polymorphic (three to nine alleles per locus). These markers will provide a system to study population segregation and diversity, gene flow, dispersal and inbreeding.     

Polymorphic microsatellite DNA markers in the penduline tit, Remiz pendulinus

To describe the exceptional mating system of the penduline tit, Remiz pendulinus, we aim to combine field observation records with DNA analysis based on polymorphic microsatellite DNA markers. Here we describe features of nine loci and their corresponding polymerase chain reaction primers. The observed number of alleles varied from four to seven and the observed heterozygosity ranged from 0.419 to 0.802. Neither of the loci is sex-linked and as linkage disequilibrium analysis showed they assort independently. Seven of the nine loci were polymorphic in the Cape penduline tit, Anthoscopus minutus.     

Chronic malaria infections increase family inequalities and reduce parental fitness: experimental evidence from a wild bird population

Avian malaria parasites (Plasmodium) occur commonly in wild birds and are an increasingly popular model system for understanding host–parasite co‐evolution. However, whether these parasites have fitness consequences for hosts in endemic areas is much debated, particularly since wild‐caught individuals almost always harbour chronic infections of very low parasite density. We used the anti‐malarial drug Malarone^TM to test experimentally for fitness effects of chronic malaria infection in a wild population of breeding blue tits (Cyanistes caeruleus). Medication caused a pronounced reduction in Plasmodium infection intensity, usually resulting in complete clearance of these parasites from the blood, as revealed by quantitative PCR. Positive effects of medication on malaria‐infected birds were found at multiple stages during breeding, with medicated females showing higher hatching success, provisioning rates and fledging success compared to controls. Most strikingly, we found that treatment of maternal malaria infections strongly altered within‐family differences, with reduced inequality in hatching probability and fledging mass within broods reared by medicated females. These within‐brood effects appear to explain higher fledging success among medicated females and are consistent with a model of parental optimism in which smaller (marginal) offspring can be successfully raised to independence if additional resources become available during the breeding attempt. Overall, these results demonstrate that chronic avian malaria infections, far from being benign, can have significant effects on host fitness and may thus constitute an important selection pressure in wild bird populations.     

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

Summary High 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.