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.     

Effects of habitat fragmentation on provisioning rates, diet and breeding success in two species of tit (great tit and blue tit)

The aim of this study was to examine the effects of forest fragmentation on the ability of parent birds to provide their young with an adequate food supply. To examine whether prey population densities of the great tit (Parus major L.) and the blue tit (P. caeruleus L.) vary between study areas in different forest size classes we compared provisioning rates and chick diet and related these parameters to breeding success. We filmed 217 nests over two breeding seasons and collected data on frass fall as a general estimate of caterpillar availability. Nests which were attended by none or one parent only during filming (n = 46) were excluded from the analyses. In both years and for both species feeding rates were highest in the smallest fragments and lowest in the large forest. There was also a suggestion that differences in feeding rates between areas vary between years. We found no consistent tendency for prey size to change with forest size, although both species brought slightly smaller prey items to the nest in the smallest forest fragments and feeding rates correlated negatively with prey size. Caterpillars were the main item fed to nestlings, in both species. We found no evidence to suggest that either frass fall or the proportion of caterpillars in the diet varied with forest size. There was also no correlation between mean frass fall and the total number of caterpillars brought to the nests, in either species. Breeding success, as measured by clutch size, brood size, fledging weight and fledging success, did not differ between the small fragments and the large forest, in either species. There was also no relationship between provisioning rate (as concerns volume of prey fed to nestlings and the quality of chick diet) and breeding success parameters. In conclusion, this study does not suggest suboptimal foraging or breeding conditions in small fragments compared to a nearby large forest, for either species. Received: 23 June 1997 / Accepted: 29 December 1997     

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     

Local weather conditions have complex effects on the growth of blue tit nestlings

Adverse weather conditions are expected to result in impaired nestling development in birds, but empirical studies have provided equivocal support for such a relationship. This may be because the negative effects of adverse weather conditions are masked by parental effects. Globally, ambient temperatures, rainfall levels and wind speeds are all expected to increase in a changing climate and so there is a need for a better understanding of the relationship between weather conditions and nestling growth. Here, we describe a correlative study that examined the relationships between local temperatures, rainfall levels and wind speeds and the growth of individual blue tit (Cyanistes caeruleus) nestlings in relation to their hatching order and sex. We found that changes in a range of morphological characters were negatively related to both temperature and wind speed, but positively related to rainfall. These patterns were further influenced by the hatching order of the nestlings but not by nestling sex. This suggests that the predicted changes in local weather conditions may have complex effects on nestling growth, but that parents may be able to mitigate the adverse effects via adaptive parental effects. We therefore conclude that local weather conditions have complex effects on avian growth and the implications for patterns of avian growth in a changing climate are discussed.     

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.     

Experimental evidence for state-dependent nest weight in the blue tit, Cyanistes caeruleus

Parental investment in reproduction is generally limited by food availability, and so avian life-history research has traditionally focused on the brood rearing phase, when food requirements are greatest. Only relatively recently has the focus extended to the incubation phase, and even more recently to the nest-building phase, where observational and comparative evidence suggest that avian nest building is an energetically expensive and time-consuming activity. We aimed to experimentally test the limitations on this cost in a hole-breeding passerine, the blue tit (Cyanistes caeruleus), by providing supplementary food to experimental pairs during the nest-building period. In comparison with control females, that did not receive supplementary food, experimental females constructed heavier nests, with greater amounts of moss base but similar amounts of cup lining, despite there being no differences in the time taken to build the nest. This study provides empirical support for the hypothesis that avian nest building is a costly behaviour, limited by food availability.     

密点麻蜥的食性及摄食的季节性变化

分析甘肃民勤２０９只成体和４３只幼体密点麻蜥胃含物表明,密点麻蜥是以动物性食物为主的杂食性动物,主要以甲虫和白刺果为食．成体出蛰后即开始强烈摄自,摄食率均在８３％以上．食性季节性变化显著,春季仅以动物为食,夏季动物性和植物性食物各半,秋季植物性食物占１／３．食物种类多样性和营养生态位宽度随体形增大而增加．     

No effect of parental quality or extrapair paternity on brood sex ratio in the blue tit (Parus caeruleus)

Sex allocation theory predicts that parents should manipulatebrood sex ratio in order to maximise the combined reproductivevalue of their progeny. Females mating with high quality malesshould, therefore, be expected to produce brood sex ratiosbiased towards sons, as male offspring would receive a relativelygreater advantage from inheritance of their father's characteristicsthan would their female siblings. Furthermore, it has been suggested that sex allocation in chicks fathered through extrapair fertilizations should also be biased towards sons. Contraryto these predictions, we found no evidence that the distributionof sex ratios in a sample of 1483 chicks from 154 broods ofblue tits (Parus caeruleus) deviated significantly from thatof a binomial distribution around an even sex ratio. In addition,we found no significant effect on brood sex ratio of the individualquality of either parent as indicated by their biometrics, feather mite loads, time of breeding, or parental survival. This suggeststhat females in our population were either unable to manipulateoffspring sex allocation or did not do so because selectionpressures were not strong enough to produce a significant shiftaway from random sex allocation. The paternity of 986 chicks from 103 broods was determined using DNA microsatellite typing.Extrapair males sired 115 chicks (11.7%) from 41 broods (39.8%).There was no significant effect of paternity (within-pair versusextrapair) on the sex of individual offspring. We suggest that,in addition to the weakness of selection pressures, the possiblemechanisms responsible for the allocation of sex may not besufficiently accurate to control offspring sex at the levelof the individual egg.     

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.     

Climatic discrimination of Mediterranean broad-leaved sclerophyllous and deciduous forests in central Spain

Abstract. Climatic differences between three types of deciduous (Quercus pyrenaica) and three types of sclerophyllous (Quercus rotundifolia) Mediterranean forests in the Spanish Sistema Central were analyzed by means of Canonical Discriminant Analysis and Jancey's Discriminant Analysis, applied in successive steps to data from 252 meteorological stations. Climatic data included temperature and precipitation records as well as bioclimatic indices. Discriminant analysis was applied to broad-leaved sclerophyllous and deciduous forest communities sampled at each meteorological station using phytosociological methods. Annual and seasonal (summer, spring) water availability are the most important factor controlling the distribution of the two physiognomic forest types; southwestern associations of Quercus pyrenaica and Q. rotundifolia differ from their colder homologues by annual and monthly temperatures; western associations were separated from eastern ones in terms of annual and seasonal precipitation gradients. Discriminant analysis was a good technique to explore climatic gradients not shown by other general ordination or classification methods.     

Caterpillar abundance in the territory affects the breeding performance of great tit Parus major minor

The effects of caterpillar food supply on the breeding performance of a population of the Japanese great tit Parus major minor were investigated. Since more than 90% of the food items in our study site were caterpillars living on trees, we estimated the food availability using 20 frass traps per hectare. The sampling error of this method was about 10% on average, which was accurate enough to detect differences between territories. Food abundance at laying in each territory affected the timing of egg laying. However, food amount after hatching was correlated with clutch size. No relationship was found between fledgling quality and food availability, probably because the effects of local variation in food abundance could be canceled out by parental effort such as extending the foraging area. There was a significant negative correlation between the length of the nestling period and food availability. We suggest that parent tits decide the timing of fledging at the point where two factors, predation risk before fledging and additional improvement of nestling quality, are balanced. Food availability just after fledging affected the length of post-fledging parental care; it seems that fledglings in “poor” territories would have had difficulty in finding food and hence needed to depend on their parents longer than those in “rich” territories. Received: 10 June 1997 / Accepted: 29 December 1997     

Feeding ecology and daily ration of the pelagic armorhead,Pseudopentaceros wheeleri,at Southeast Hancock Seamount

Synopsis The unusual life history of pelagic armorhead, Pseudopentaceros wheeleri, includes occupation of epipelagic subarctic and transitional waters of the northeast Pacific Ocean and Gulf of Alaska as subadults and subsequent recruitment to the southern Emperor and northern Hawaiian Ridge (SE-NHR) seamounts in the central North Pacific Ocean where they assume a demersal existence and begin maturation as adults. Feeding ecology and daily ration were examined in pelagic armorhead during field studies conducted at Southeast Hancock Seamont (29°48′ N,179°04′ E) in the SE-NHR seamounts. A total of 1133 stomachs collected during June-July 1985, January 1988, and July 1988 were examined for contents; only 19.7% contained food items. The diet consisted of pelagic tunicates and other micronektonic animals that are associated with the vertically migrating sonic scattering layer that is advected over the seamount from the adjacent oceanic environment. Peak feeding occurred in the early morning on migrating organisms likely trapped by the expanse of the seamount summit during the morning descent. Newly recruited armorhead had lower feeding rates than older, previously recruited fish. Estimates of daily ration for previously recruited armorhead were 1.17 g (0.26% of body weight) in the summer of 1985 and 12.87 g (2.86% of body weight) in the summer of 1988. Since estimated values for the evacuation coefficient and the duration of the daily feeding period did not differ between sampling periods, the increase in daily ration was due to an increase in feeding rate. The proportion of the diet composed of pelagic tunicates was also higher in 1988.     

Identification of hypervariable single locus minisatellite DNA probes in the blue tit Parus caeruleus

We report the isolation of a set of hypervariable minisatellite DNA sequences from a blue tit Parus caeruleus genomic DNA library. In our strategy, we cloned a minisatellite-rich DNA fraction into a charomid vector. The resulting cosmid library was screened with the two minisatellite DNA probes 33.6 and 33.15 for recombinants containing a minisatellite DNA insert. A total of 233 positive clones were isolated. Of 37 clones that have been analysed, nine gave polymorphic signals and can be used as single locus probes (SLPs). Four of the SLPs were investigated in more detail. The number of alleles, the heterozygosity and the mutation rate were estimated. Linkage analysis revealed that two of these loci were linked. The SLPs are of value to studies of the mating system and reproductive success in the blue tit, and may also be useful in population genetic studies.     

渤海南部牙鲆的食性及摄食的季节性变化

1051尾渤海南部牙鲆的胃含物分析表明,渤海南部牙鲆为捕食性(游泳生物食性)鱼类,其食物成分有30余种,以底栖虾类中的口虾蛄、鲜明鼓虾、日本鼓虾、脊尾褐虾,软体动物的日本枪乌贼及鱼类中的斑(鱼祭)、鳀鱼、黄鲫、六丝矛尾虾虎鱼及焦氏舌鳎等动物为主要食物。牙鲆除冬季外,其余时间均强烈摄食,而且其摄食的季节性变化明显:春季以鱼类(重量占80.9％)及甲壳类(18.6％)为主要食物;夏季以鱼类(83.5％)及软体动物类(13.1％)为主要食物;秋季主要摄食鱼类(87.9％);冬季则主要摄食鱼类(73.2％)及甲壳类(19.7％)。     

Seasonal variation in reproductive success and post-nuptial moult of blue tits in southern Europe: an experimental study

Post-nuptial moult and reproductive success were studied in relation to timing of breeding in blue tits, Parus caeruleus, breeding in southern Europe. A group of experimentally delayed pairs was created by removing first clutches, thereby inducing late repeat clutches. Reproductive success and post-nuptial moult of delayed pairs were compared with both control pairs that bred early and unmanipulated late-breeding pairs. Delayed pairs fledged fewer young and with a lower body mass than control pairs. However, the number of fledged young and fledgling mass did not differ between delayed and late-breeding pairs. These results were more consistent with the date hypothesis, and it is concluded that the timing of breeding and reproductive success may be causally related in the blue tit. This study reveals a harmful effect of relaying on female body mass at the end of the nestling period. Therefore, females apparently pay the costs of relaying, since a reduction in body mass during the nestling period may be accompanied by a lowered survival probability. Delayed and late-breeding males often began moulting while still feeding young, but neither control males nor females from the three study groups did so. These results support the view that timing-related energy constraints on breeding may be important causes of a seasonal decline in reproductive success at different latitudes. Received: 15 March 1999 / Accepted: 19 July 1999     

Diversity of feeding habitats and diet composition in the turtle doves Streptopelia turtur to buffer loss and modification of natural habitats during breeding season

The approach of the birds to use physical and alimentary resources in degraded and modified natural habitats is an important aspect of their adaptation. This study was undertaken during 2016-2017 at forty habitats in the Moulouya plain, Morocco to examine behavioral diet, habitat use and foraging ecology of turtle dove, Streptopelia turtur. We monitored turtle doves in four major plots namely cereal plots, lucerne farms, apple orchards, and stations in the Ansgmir River covering 40 habitats. Digestive tract contents were also identified and evaluated for 68 Turtle Doves shot by hunters during two consecutive years. The results showed that the turtle doves use a variability of feeding habitats. The cereal cultivation seemed to be more preferable habitat for feeding especially in the month of May, the first breeding phase of the year. But, during the months of August and July, the riverbanks were the preferred habitat for turtle doves. The diet of this species is polyphagous and diverse with a granivorous tendency. Diet analysis showed that soft wheat and barley seeds constituted 44.53% and 38.74% respectively followed by barley seeds with 38.74% and sand stones (9.16%) of the seeds eaten by Turtle Doves. However, moderate proportion of elements (7.32%) remained undefined. All these aspects, including the variability of feeding habitats and the wide diet seem to be an adaptive strategy followed by turtle dove to counter the degradation and the modification of its natural feeding habitats.     

Evolution of sexual size dimorphism in birds: test of hypotheses using blue tits in contrasted Mediterranean habitats

Many hypotheses, either sex‐related or environment‐related, have been proposed to explain sexual size dimorphism in birds. Two populations of blue tits provide an interesting case study for testing these hypotheses because they live in contrasting environments in continental France and in Corsica and exhibit different degree of sexual size dimorphism. Contrary to several predictions, the insular population is less dimorphic than the continental one but neither the sexual selection hypothesis nor the niche variation hypothesis explain the observed patterns. In the mainland population it is advantageous for both sexes to be large, and males are larger than females. In Corsica, however, reproductive success was greater for pairs in which the male was relatively small, i.e. pairs in which sexual size dimorphism is reduced. The most likely explanation is that interpopulation differences in sexual size dimorphism are determined not by sex‐related factors, but by differences in sex‐specific reproductive roles and responses to environmental factors. Because of environmental stress on the island as a result of food shortage and high parasite infestations, the share of parents in caring for young favours small size in males so that a reduced sexual size dimorphism is not the target of selection but a by‐product of mechanisms that operate at the level of individual sexes.