Social stability and daily body mass gain in great tits

This paper presents the results of an experiment on the influenceof changes in flock composition and social rank on the patternof daily mass gain in captive wintering great tits. We createdflocks of three male great tits and scored dominance when rankshad stabilized after a few days. We then moved the dominantbird from each flock into a neighboring flock, thus creatinga certain amount of social instability and unpredictable resourceaccess. Both before and after moving the dominants, we collectedindividual body mass data three times a day. When the hierarchieshad stabilized in the second flocks, individual ranks were scored.After the dominant birds were moved, the rate of daily massgain increased for the average individual. There was also adecrease in average morning body mass, so the birds startedthe day at a lower mass and ended at a slightly higher massthan previously. Dominants were on average heavier than subordinates,but we found no statistically significant effect of rank changeon body mass. The pattern of daily mass gain was linear overtime both before and after treatment. We conclude that an impressionof unpredictable resource access can be created by disturbingthe social structure in a flock, with similar effect as fromrestricting individual access to feeders. Within a flock, instabilityof the social hierarchy thus seems to have an influence on bodymass management in addition to that of environmental stochasticity.     

Early learning affects social dominance: interspecifically cross-fostered tits become subdominant

Social dominance influences the outcome of competitive interactionsover limited resources, and may hence be important for individualfitness. Theory thus predicts that its heritability will below and that non-genetic determinants of dominance should prevail.In this field experiment we reciprocally cross-fostered greattits (Parus major) to blue tits (Parus caeruleus) to investigatethe impact of early social experience on dominance status incompetition over food during winter. Controlling for potentialeffects of age, size, sex and site-related dominance, we showthat cross-fostered birds of both species were subdominant toconspecific immigrants, while controls originating from unmanipulatedbroods were dominant to conspecific immigrants. Furthermore,blue tits reared by blue tit parents but with at least one greattit broodmate had lower dominance status relative to conspecificimmigrants than did controls. Although great tits generallydominated blue tits, cross-fostered birds of both species initiatedmarginally more fights against the other species than did theirrespective controls, suggesting faulty species recognition.Since both social parents and broodmates strongly influencethe dominance behavior of offspring later in life, we concludethat social conditions experienced at an early age are crucialfor the determination of subsequent social dominance.     

The influence of predation risk on threat display in great tits

In wintering birds, conflicts over food are often resolved bythreat displays. For displays to be effective, there oughtto be a cost associated with displaying. We investigated whetherincreased vulnerability to predators due to reduced vigilancecould be such a cost. Conflicts ought then to be resolved usingfewer or less intense displays in conditions of high risk. We also looked for differences between dominants and subordinatesin their reaction to risk. Because there is considerable evidencethat subordinate wintering birds forage in riskier places thandominants, one might expect dominants to be less successfulin conflicts under high predation risk. In our experiment,nine flocks of four or five wintering male great tits were keptin outdoor aviaries. In the predation risk treatment, a stuffedpygmy owl was briefly shown before birds were allowed accessto a feeder. In the control treatment the owl did not appear.The predator presentation caused a reduction in the amountof aggression shown by subordinates, whereas for dominants there was no statistically significant change. Dominants were at leastas successful in subduing subordinates under high risk as underlow risk. A possible interpretation is that our experimentreflected a natural foraging situation for great tits, whereephemeral resources can appear unpredictably. In such situations,dominants may need to be bold to gain priority of access even under increased risk of predation, whereas a subordinate wouldgain little by risking a conflict with small chances of winning.     

Context dependence of personalities: risk-taking behavior in a social and a nonsocial situation

Individuals of many species differ consistently in their behavioralreaction to mild novel challenges. Suites of these behaviorsare referred to as behavioral syndromes or personalities. Personalitytraits are often phenotypically and genetically correlated.Therefore, animal personalities are generally considered asbroad characteristics, with one underlying genetical and physiologicalmechanism that is expressed across situations and contexts.Because there are carryover effects between situations, animalsare not entirely flexible in their behavior in each situation.This may cause behaviors to seem nonadaptive in isolated situations.To test whether individuals with different personalities couldreact differently to changes in their environment, we studiedcontext dependence of personalities in the great tit (Parusmajor). We tested birds categorized as either fast or slow explorersfor their latency to come back to a feeding table after a mildstartle (risk-taking behavior) in a nonsocial followed by asocial context. We found that the relation between exploratorybehavior and risk-taking behavior depended on the social context.Females in general returned later in the social test, whilemale reaction to the presence of a conspecific was dependenton their behavioral type. Slow males thereby reacted to thebehavior of the companion and fast males did not. These resultsshow that although personalities have a rigid structure therelation between personality traits is context dependent. Theseresults are discussed in the perspective of the adaptive significanceand maintenance of personalities.     

The function of extrapair paternity in blue tits and great tits: good genes or fertility insurance?

DNA fingerprinting of an island population of blue tits andgreat tits in southeast Norway revealed that extrapair paternityaccounted for 36% (17/47) and 27% (15/55) of broods and for7% (31/466) and 8% (33/408) of young in the two species, respectively.Cuckolded males did not differ from noncuckolded males withrespect to morphology, age, or survival. There was no seasonalpattern in the frequency of extrapair paternity, and males showedno individual consistency in paternity loss over multiple broods.Extrapair offspring did not grow faster, they did not fledgewith a higher body mass, and they did not show a higher localsurvival rate than their half siblings. Hence, there was noevidence of any association between extrapair paternity andmale phenotypic or genotypic quality. Extrapair offspring wererandomly distributed among broods, with the only exceptionsof one blue tit and two great tit broods in which all young(six to nine) were sired by an extrapair male. This patternis best explained by a small proportion of males (2%–4%)being infertile and by most females performing a few extrapaircopulations as insurance against laying infertile eggs. We concludethat the results suggest a role for fertility insurance butthat alternative functional explanations to extrapair paternityin these populations cannot yet be ruled out.     

Clutch size and malarial parasites in female great tits

life-history models predict an evolutionary trade-off in theallocation of resources to current versus future reproduction.This corresponds, at the physiological level, to a trade-offin the allocation of resources to current reproduction or tothe immune system, which will enhance survival and thereforefuture reproduction. For clutch size, life-history models predicta positive correlation between current investment in eggs andthe subsequent parasite load. In a population of great tits,we analyzed the correlation between natural clutch size of femalesand the subsequent prevalence of Plasmodium spp., a potentiallyharmful blood parasite. Females that showed, 14 days after hatchingof the nestlings, an infection with Plasmodium had a significantlylarger clutch (9.3 eggs ± 0.5 SE, n = 18) than uninfectedfemales (8.0 eggs ± 0.2 SE, n = 80), as predicted bythe allocation trade-off. Clutch size was positively correlatedwith the prevalence of Plasmodium, but brood size 14 days afterhatching was not. This suggests that females incur higher costsduring laying the clutch than during rearing nestlings. Infectionstatus of some females changed between years, and these changeswere significantly correlated with a change in clutch size aspredicted by the trade-off. The link between reproductive effortand parasitism may represent a possible mechanism by which thecost of egg production is mediated into future survival andmay thereby be an important selective force in the shaping ofclutch size     

Health impact of blood parasites in breeding great tits

Hypotheses of hemoparasite-mediated sexual selection and reproductive costs rely on the assumption that avian blood parasite infections are harmful to their hosts. To test the validity of this assumption, we examined the health impact of Haemoproteus blood parasites on their great tit (Parus major) host. We hypothesised that if blood parasites impose any serious health impact on their avian hosts, then infected individuals must differ from uninfected ones in respect to hemato-serological general health and immune parameters. A 3-year study of two great tit populations, breeding in contrasting (urban and rural) habitats in south-east Estonia, revealed that Haemoproteus blood parasites affected the health state of their avian hosts. Infected individuals had elevated lymphocyte hemoconcentration and plasma gamma-globulin levels, indicating that both cell-mediated and humoral immune response mechanisms are involved in host defence. The effect of parasites on cell-mediated immunity was both age- and sex-specific, as infection status affected peripheral blood lymphocyte counts only in males, and among these, the magnitude of response was greater in old individuals than yearlings. Heterophile hemoconcentration and plasma albumin levels were not affected by infection status, suggesting that blood stages of Haemoproteus infection do not cause a severe inflammatory response. Parasitism was not related to hematocrit values, indicating that Haemoproteus infection does not cause anemia. In two years, infected individuals were heavier than uninfected ones in the urban but not in the rural study area. This suggests, that under certain circumstances (possibly related to reproductive tactics), breeding great tits may avoid losing body mass in order to save resources for an anti-parasite immune response. Received: 16 February 1998 / Accepted: 22 May 1998     

Association between DRD4 gene polymorphism and personality variation in great tits: a test across four wild populations

Polymorphisms in the dopamine receptor D4 gene (DRD4) have been related to individual variation in novelty‐seeking or exploratory behaviour in a variety of animals, including humans. Recently, the human DRD4 orthologue was sequenced in a wild bird, the great tit (Parus major) and a single nucleotide polymorphism in exon 3 of this gene (SNP830) was shown to be associated with variation in exploratory behaviour of lab‐raised individuals originating from a single wild population. Here we test the generality of this finding in a large sample of free‐living individuals from four European great tit populations, including the originally sampled population. We demonstrate that the association between SNP830 genotype and exploratory behaviour also exists in free‐living birds from the original population. However, in the other three populations we found only limited evidence for an association: in two populations the association appeared absent; while in one there was a nonsignificant tendency. We could not confirm a previously demonstrated interaction with another DRD4 polymorphism, a 15 bp indel in the promoter region (ID15). As yet unknown differences in genetic or environmental background could explain why the same genetic polymorphism (SNP830) has a substantial effect on exploratory behaviour in one population, explaining 4.5–5.8% of the total variance—a large effect for a single gene influencing a complex behavioural trait—but not in three others. The confirmation of an association between SNP830 genotype and personality‐related behaviour in a wild bird population warrants further research into potential fitness effects of the polymorphism, while also the population differences in the strength of the association deserve further investigation. Another important future challenge is the identification of additional loci influencing avian personality traits in the wild.     

Mhc‐linked survival and lifetime reproductive success in a wild population of great tits

Major histocompatibility complex (Mhc) genes are frequently used as a model for adaptive genetic diversity. Although associations between Mhc and disease resistance are frequently documented, little is known about the fitness consequences of Mhc variation in wild populations. Further, most work to date has involved testing associations between Mhc genotypes and fitness components. However, the functional diversity of the Mhc, and hence the mechanism by which selection on Mhc acts, depends on how genotypes map to the functional properties of Mhc molecules. Here, we test three hypotheses that relate Mhc diversity to fitness: (i) the maximal diversity hypothesis, (ii) the optimal diversity hypothesis and (iii) effect of specific Mhc types. We combine mark–recapture methods with analysis of long‐term breeding data to investigate the effects of Mhc class I functional diversity (Mhc supertypes) on individual fitness in a wild great tit (Parus major) population. We found that the presence of three different Mhc supertypes was associated with three different components of individual fitness: survival, annual recruitment and lifetime reproductive success (LRS). Great tits possessing Mhc supertype 3 experienced higher survival rates than those that did not, whereas individuals with Mhc supertype 6 experienced higher LRS and were more likely to recruit offspring each year. Conversely, great tits that possessed Mhc supertype 5 had reduced LRS. We found no evidence for a selective advantage of Mhc diversity, in terms of either maximal or optimal supertype diversity. Our results support the suggestion that specific Mhc types are an important determinant of individual fitness.     

Experimental evidence for density dependence of reproduction in great tits

1. Density dependence of avian reproduction has often been analysed using correlations between annual mean reproductive output and population density. Experiments are necessary to prove that density is the cause of the observed patterns, but so far, three out of four experimental studies do not support a direct causal effect of density on reproduction.
2. This paper presents experimental evidence that reproductive decisions in great tits, Parus major L., are causally affected by breeding density. The breeding density of great tits was manipulated by providing nest-boxes at different densities in an ecologically homogeneous area.
3. Within years the densities in the high and low density plots differed approximately 8-fold. During the 11 years of the experiment, clutch size, nestling mass and the proportion of birds starting a second brood were all lower in the high density plot. In 5 years with equal breeding densities in both parts, clutch size did not differ between the plots. The patterns found were consistent with the density effects as predicted from the non-experimental data.     

Climate change and breeding parameters of great and blue tits throughout the western Palaearctic

Increasing evidence suggests that climate change has consequences on avian breeding phenology. Here, variations in laying date and clutch size of great tit Parus major and blue tit Parus caeruleus within and between breeding populations through the western Palaearctic are examined in relation to climatic fluctuations, measured by the winter North Atlantic Oscillation (NAO) index. Within and across breeding sites, laying date was related to winter‐NAO index such that great and blue tit females lay earlier after warmer, moister winters (positive values of winter NAO‐index). The present study shows that for most populations there is an advancement of laying date, but the rate of change with respect to NAO significantly differed geographically across the western Palaearctic and did not differ between species. However, clutch size of great and blue tits was not affected by climatic fluctuations, presumably because the whole season is being shifted, but not in relation to food supplies. These combined analyses for the two species controlled for potentially confounding variables such as latitude, longitude, elevation and habitat of each study site.     

Gene flow and immigration rate in an island population of great tits

It is generally recognized that immigration and gene flow cannot be equated, but few detailed quantitative comparisons of these processes have been made over the entire lifetime of individual animals. We analyzed data collected in a longterm study of an island population of great tits Parus major, and tested two assumptions frequently made in population genetic studies. (1) The assumption that there is no difference in reproductive output between immigrant and resident breeding birds was refuted for females but not for males. Lifetime production of local recruits (LRS) was reduced by 37% in immigrant females, because female immigrants tended to leave the island after breeding, and thus reproduced for fewer years. Female LRS was density dependent, but the effect of density was independent of status (resident/immigrant). (2) The assumption that mating was random with respect to status was also refuted: assortative mating was found, even when temporal and spatial aggregation of immigrants was controlled for. Thus both assumptions were refuted, and gene flow was lower than immigration rate.     

No evidence for MHC class I‐based disassortative mating in a wild population of great tits

Genes of the major histocompatibility complex (MHC) are regarded as a potentially important target of mate choice due to the fitness benefits that may be conferred to the offspring. According to the complementary genes hypothesis, females mate with MHC dissimilar males to enhance the immunocompetence of their offspring or to avoid inbreeding depression. Here, we investigate whether selection favours a preference for maximally dissimilar or optimally dissimilar MHC class I types, based on MHC genotypes, average amino acid distances and the functional properties of the antigen‐binding sites (MHC supertypes); and whether MHC type dissimilarity predicts relatedness between mates in a wild great tit population. In particular, we explore the role that MHC class I plays in female mate choice decisions while controlling for relatedness and spatial population structure, and examine the reproductive fitness consequences of MHC compatibility between mates. We find no evidence for the hypotheses that females select mates on the basis of either maximal or optimal MHC class I dissimilarity. A weak correlation between MHC supertype sharing and relatedness suggests that MHC dissimilarity at functional variants may not provide an effective index of relatedness. Moreover, the reproductive success of pairs did not vary with MHC dissimilarity. Our results provide no support for the suggestion that selection favours, or that mate choice realizes, a preference for complimentary MHC types.     

Repeatability and heritability of exploratory behaviour in great tits from the wild

We investigated whether individual great tits, Parus major, vary consistently in their exploratory behaviour in a novel environment and measured the repeatability and heritability of this trait. Wild birds were caught in their natural habitat, tested in the laboratory in an open field test on the following morning, then released at the capture site. We measured individual consistency of exploratory behaviour for recaptured individuals (repeatability) and estimated the heritability with parent-offspring regressions and sibling analyses. Measures of exploratory behaviour of individuals at repeated captures were consistent in both sexes and study areas (repeatabilities ranged from 0.27 to 0.48). Exploration scores did not differ between the sexes, and were unrelated to age, condition at fledging or condition during measurement. Heritability estimates were 0.22-0.41 (parent-offspring regressions) and 0.37-0.40 (sibling analyses). We conclude that (1) consistent individual variation in open field behaviour exists in individuals from the wild, and (2) this behavioural variation is heritable. This is one of the first studies showing heritable variation in a behavioural trait in animals from the wild, and poses the question of how this variation is maintained under natural conditions. Copyright 2002 The Association for the Study of Animal Behaviour. Published by Elsevier Science Ltd. All rights reserved.     

Brood size, sibling competition, and the cost of begging in great tits (Parus major)

Evolutionary theory of parent-offspring conflict explains beggingdisplays of nestling birds as selfish attempts to influenceparental food allocation. Models predict that this conflictmay be resolved by honest signaling of offspring need to parents,or by competition among nestmates, leading to escalated beggingscrambles. Although the former type of models has been qualitativelysupported by experimental studies, the potential for a beggingcomponent driven by scramble competition cannot be excludedby the evidence. In a brood-size manipulation experiment withgreat tits, Parus major, we explored the scramble componentin the begging activity of great tit nestlings by investigatingthe mechanisms of sibling competition in relation to brood size.While under full parental compensation, the feeding rate pernestling will remain constant over all brood sizes for bothtypes of models; the scramble begging models alone predict anincrease in begging intensity with brood size, if begging costsdo not arise exclusively through predation. Great tit parentsadjusted feeding rates to brood size and fed nestlings at similarrates and with similar prey sizes in all three brood-size categories.Despite full parental compensation, the begging and food solicitationactivities increased with experimental brood size, whereas nestlingbody condition deteriorated. These findings support a scramblecomponent in begging and suggest that the competition-inducedcosts of food solicitation behavior play an important role inthe evolution of parent-offspring communication.     

The cheek plumage patch is an amplifier of dominance in great tits

Amplifiers are signals that enhance the perception of other signals or cues, but no studies to date provide empirical evidence for the role of these signals in a reproductive context. Here we use the white cheek patch of great tits as a model for studying this issue. Aggressive interactions decrease patch immaculateness, so patch size may be an amplifier of dominance, that is, more clearly reveal status. If so, in high-quality individuals patch size should correlate positively with reproductive success (here estimated by laying date, assuming that the earlier the better), whereas low-quality individuals with a large patch should only more clearly reveal their low quality and thus suffer low reproductive success, which is exactly the pattern found in males. In contrast, the cheek patch does not seem to function as an amplifier in female great tits.     

A behavioural syndrome in the field cricket Gryllus integer: intrasexual aggression is correlated with activity in a novel environment

Behavioural syndromes, or suites of correlated behaviours across different contexts and situations, have recently drawn attention from evolutionary biologists. In the field cricket Gryllus integer , males are aggressive with one another and fight vigorously over females and territories. We examined whether aggressiveness with other males was correlated with activity in a potentially dangerous context (a novel environment) in laboratory-raised virgin males. Aggressiveness was measured as fighting ability against a weight-matched opponent. First, we measured each cricket's latency to become active in a novel environment and latency to emerge from a refuge within a novel environment. Next, we determined which of two weight-matched males was more aggressive, by pitting the males together in an agonistic contest and counting the number of fights won by each male. More aggressive males, who won more fights, had shorter latencies to become active when placed in a novel environment and shorter latencies to emerge from a safe refuge. These results suggest that a behavioural syndrome exists in G. integer , in which more aggressive males are also more active in general, and possibly less cautious towards predation risk.  © 2007 The Linnean Society of London, Biological Journal of the Linnean Society , 2007, 91 , 475–482.