The Tradeoff Between Territorial Defense and Foraging in the Great Tit (Parus major)

We analyze territorial behavior in terms of decisions abouttime allocation.Such decisions must be made whenever time investedin territorial defense cannot be devoted to feeding, and viceversa. We describe the ecology and territorial behavior of thegreat tit (Parus major) to show that a tradeoff exists, andthen outline a series of laboratory and field experiments inwhich the value of feeding or defense was experimentally manipulated.Territorialmale great tits began to invest more heavily in territorialvigilance after encountering intruders, but the increase invigilance depended on the rate at which they could feed, aswell as their hunger level. We outline a dynamic analysis thattakes account of the fact that the optimal tradeoff will changeas hunger is reduced. The results of an experimental test ofthis dynamic model are also presented. We briefly review othertechniques whereby territorial tradeoffs have been investigated.     

Intraspecific Variation and Interspecific Differences in the Bacterial and Fungal Assemblages of Blue Tit (Cyanistes caeruleus) and Great Tit (Parus major) Nests

Although interest in the relationship between birds and microorganisms is increasing, few studies have compared nest microbial assemblages in wild passerines to determine variation within and between species. Culturing microorganisms from blue tit (Cyanistes caeruleus) and great tit (Parus major) nests from the same study site demonstrated diverse microbial communities with 32 bacterial and 13 fungal species being isolated. Dominant bacteria were Pseudomonas fluorescens, Pseudomonas putida, and Staphylococcus hyicus. Also common in the nests were the keratinolytic bacteria Pseudomonas stutzeri and Bacillus subtilis. Dominant fungi were Cladosporium herbarum and Epicoccum purpurascens. Aspergillus flavous, Microsporum gallinae, and Candida albicans (causative agents of avian aspergillosis, favus, and candidiasis, respectively) were present in 30%, 25%, and 10% of nests, respectively. Although there were no differences in nest mass or materials, bacterial (but not fungal) loads were significantly higher in blue tit nests. Microbial species also differed interspecifically. As regards potential pathogens, the prevalence of Enterobacter cloacae was higher in blue tit nests, while Pseudomonas aeruginosa—present in 30% of blue tit nests—was absent from great tit nests. The allergenic fungus Cladosporium cladosporioides was both more prevalent and abundant in great tit nests. Using discriminant function analysis (DFA), nests were classified to avian species with 100% accuracy using the complete microbial community. Partial DFA models were created using a reduced number of variables and compared using Akaike’s information criterion on the basis of model fit and parsimony. The best models classified unknown nests with 72.5–95% accuracy using a small subset of microbes (n = 1–8), which always included Pseudomonas agarici. This suggests that despite substantial intraspecific variation in nest microflora, there are significant interspecific differences—both in terms of individual microbes and the overall microbial community—even when host species are closely related, ecologically similar, sympatric, and construct very similar nests.     

Foraging Behaviour of Subordinate Great Tits (Parus major). Can Morphology Reduce the Cost of Subordination?

This paper studies the magnitude of the behavioural shift, from forage standing to forage hanging, of subordinate great tits ( Parus major ) in two different social contexts: feeding solitarily vs. feeding with a dominant conspecific. The aim is to test the hypothesis that differences in morphological design provide subordinates with varying abilities to reduce the presumed costs of subordination. We find that different subordinate individuals change the foraging behaviour, occupying a different niche when an intra-specific competitor is present. Morphology linked to sexual dimorphism, specifically body mass, is the factor responsible for the different magnitudes of change. Lighter subordinates can remain longer than heavier ones at the feeding patch without interrupting their foraging. Thereby, the former reduce the costs of being subordinate more than the latter. Among subordinates, females are lighter than males; they also spend more time feeding in the presence of a dominant conspecific than males do. No differences are found between age categories. We find no relationship between tarsus length and individual ecological plasticity. Our results support the idea that the ecological plasticity due to morphological differences is a mechanism that allows subordinate individuals to overcome costs associated with subordination.     

Greenhouse Studies on the Effect of Marigolds (Tagetes spp.) on Four Meloidogyne Species

The effects of preplanted marigold on tomato root galling and multiplication of Meloidogyne incognita, M. javanica, M. arenaria, and M. hapla were studied. Marigold cultivars of Tagetes patula, T. erecta, T. signata, and a Tagetes hybrid all reduced galling and numbers of second-stage juveniles in subsequent tomato compared to the tomato-tomato control. All four Meloidogyne spp. reproduced on T. signata ''Tangerine Gem''. Several cultivars of T. patula and T. erecta suppressed galling and reproduction of Meloidogyne spp. on tomato to levels lower than or comparable to a fallow control. Phytotoxic effects of marigold on tomato were not observed. Several of the tested marigold cultivars are ready for full-scale field evaluation against Meloidogyne spp.     

Personality and Nest Defence in the Great Tit (Parus major)

A growing number of studies have shown that individuals differ consistently in a suite of correlated behavioural traits across various contexts and situations. Yet, most work on animal personalities has been performed under laboratory conditions and still little is known about the ecological significance of differences in personality in the wild, and the behavioural mechanisms underlying possible fitness consequences. In this study, we investigated individual differences in personality in relation to nest defence behaviour in wild great tits. Nest defence is an important aspect of parental care and involves a trade‐off between two fitness components (i.e. survival and reproduction). As a measure of personality we used exploratory behaviour in a novel environment as this has been shown to be correlated with several other behavioural traits including risk‐taking and aggression, two important behavioural components of nest defence. We found that the intensity of alarm calling towards a human intruder was positively associated with exploratory behaviour, while there was a negative association between exploration score and number of movements during nest defence. Thus, fast explorers are shown to respond more boldly towards predators in the field. More generally we show that individuals with different personalities vary in their anti‐predator and reproductive investment strategies.     

Vigilance of a Nest-building Female Blue Tit (Parus caeruleus)

Observations of a single nest-building female blue tit (Parus caeruleus) supported the hypothesis that increasing proximity of domestic cats (Felis domesticus) to the nest box increases time spent scanning the nest site area. A hypothesised decrease in time in the nest box with increased cat proximity was not confirmed. A reason for this is forwarded, and differences in time spent in/and in the vicinity of the nest box reflecting position of the nest visit in a bout, and threat posed by individual cats are discussed.     

Mixed species flocking of tits (Parus spp.): a field experiment

Summary We tested two general models of flocking behaviour, namely the antipredation model and foraging efficiency model on mixed-species tit flocks (Parus spp.). After food addition the size of mixed-species flocks was significantly less than in the control samples. In the presence of extra food significantly more birds were observed either in monospecific flocks or solitary, than during the control observations. In the presence of a living predator the birds foraged in larger mixed-specifies flocks than during the control observations. In addition, the social behaviour of Great Spotted Woodpecker, Middle Spotted Woodpecker and Nuthatch shifted to mixed-specific flocking. The size of monospecific flocks was independent of both treatments. The density of birds increased significantly after food addition, while in the predator presence the birds tended to leave the forest. These results support the view that both the antipredation model and foraging efficiency model seem to be valid for mixed-species flocking. However, in the case of monospecific flocks, the territory maintenance could be the most important factor.     

Characterizing the mammalian hair present in Great Tit (Parus major) nests

Capsule Many birds are known to incorporate mammal hair into their nest lining, but the frequency with which they use hair from different mammals is unknown. We performed one of the first detailed examinations of mammalian hair from 54 Great Tit nests. We identified 5317 hairs belonging to 21 mammal species. Almost all of the examined nests contained hair from Roe Deer (Capreolus capreolus), but hair from Wild Boars (Sus scrofa), European Hares (Lepus europaeus) and Red Foxes (Vulpes vulpes) was also common.     

Effects of population growth on Crested Tit Parus cristatus post-fledging settlement

From 1987 to 1993, the winter density of Crested Tits Parus cristatus in north Belgium increased 2.76 times. The increase was the result of high annual survival rates of both adult and first-year birds despite a low reproductive success (number of fledglings per female) due to high predation of nestlings. A fc-factor analysis assuming intermediate costs of post-fledging dispersal and settlement gave the best fit. First-year birds had two options when trying to settle in a prospective breeding territory: (1) establish a new territory during August or early September and defend it against neighbouring flocks during winter or immigrants in early spring or both, or (2) overwinter as a subordinate in a mixed-age flock and pair with a widowed adult or replace a territory owner in early spring ("hopeful dominant stategy"). Settlement strategies varied with density; at low density, only the hopeful dominant strategy was observed, whereas at higher density, both strategies occurred.     

Paraphyly of the Blue Tit (Parus caeruleus) suggested from cytochrome b sequences

The phylogenetic relationships of the Blue Tit-Azure Tit assemblage (genus Parus; Aves: Passeriformes) were studied using mitochondrial DNA sequences of 24 specimens representing seven subspecies from Eurasia and North Africa. Previous work based on comparative morphological and acoustic data suggested a division of the Blue Tit (Parus caeruleus) into two species. Our analyses clearly indicate that the Blue Tit represents a paraphyletic assemblage, including a European/Middle Asian clade that is the sister group to the Azure Tit (Parus cyanus) and a North African clade. The North African clade (teneriffae subspecies group) is a sister group to the European Blue Tit/Azure Tit clade. We suggest a division of the Blue Tit into two separate species, Eurasian Blue Tit (Parus caeruleus s. str.) and African Blue Tit (Parus teneriffae). However, our data give no support for assigning species rank to Parus cyanus flavipectus, a subspecies of the Azure Tit, as suggested by several authors on morphological grounds.     

Parasites as mediators of heterozygosity-fitness correlations in the Great Tit (Parus major)

Positive correlations between heterozygosity and fitness traits are frequently observed, and it has been hypothesized, but rarely tested experimentally, that parasites play a key role in mediating the heterozygosity-fitness association. We evaluated this hypothesis in a wild great tit (Parus major) population by testing the prediction that the heterozygosity-fitness association would appear in broods experimentally infested with a common ectoparasite, but not in parasite-free broods. We simultaneously assessed the effects of parental and offspring heterozygosity on nestling growth and found that body mass of nestlings close to independence, which is a strong predictor of post-fledging survival, increased significantly with nestling levels of heterozygosity in experimentally infested nests, but not in parasite-free nests. Heterozygosity level of the fathers also showed a significant positive correlation with offspring body mass under an experimental parasite load, whereas there was no correlation with the mothers' level of heterozygosity. Thus, our results indicate a key role for parasites as mediators of the heterozygosity-fitness correlations.     

Clutch Defence in Great Tit (Parus major) Pairs and the Concorde Fallacy

We conducted an experiment to test whether great tits (Parus major) base their decisions of clutch defence on past investment or future benefits. Results were obtained by manipulation of future benefits. Great tit pairs of an experimental group with reduced clutch-size and thus diminished benefits defended their offspring against a live raptor significantly less than a comparable, non-manipulated control group with the same amount of past investment (clutch-size, incubation stage, time of year). While in the females the difference between the two groups was obvious from the beginning of a trial, the difference in the males developed only during the course of a trial, suggesting that the male bases its response on the female's response deficit. A more onerous explanation of the female signaling the male the egg loss by using a language-like symbol is not supported by the data. The experiment thereby permits the conclusion that at least the great tit female avoids committing the Concorde fallacy in the strict sense.     

Plant Species Composition Alters the Sign and Strength of an Emergent Multi-Predator Effect by Modifying Predator Foraging Behaviour

The prediction of pest-control functioning by multi-predator communities is hindered by the non-additive nature of species functioning. Such non-additivity, commonly termed an emergent multi-predator effect, is known to be affected by elements of the ecological context, such as the structure and composition of vegetation, in addition to the traits of the predators themselves. Here we report mesocosm experiments designed to test the influence of plant density and species composition (wheat monoculture or wheat and faba bean polyculture) on the emergence of multi-predator effects between Adalia bipunctata and Chrysoperla carnea, in their suppression of populations of the aphid Metopolophium dirhodum. The mesocosm experiments were followed by a series of behavioural observations designed to identify how interactions among predators are modified by plant species composition and whether these effects are consistent with the observed influence of plant species composition on aphid population suppression. Although plant density was shown to have no influence on the multi-predator effect on aphid population growth, plant composition had a marked effect. In wheat monoculture, Adalia and Chrysoperla mixed treatments caused greater suppression of M. dirhodum populations than expected. However this positive emergent effect was reversed to a negative multi-predator effect in wheat and faba bean polyculture. The behavioural observations revealed that although dominant individuals did not respond to the presence of faba bean plants, the behaviour of sub-dominants was affected markedly, consistent with their foraging for extra-floral nectar produced by the faba bean. This interaction between plant composition and predator community composition on the foraging behaviour of sub-dominants is thought to underlie the observed effect of plant composition on the multi-predator effect. Thus, the emergence of multi-predator effects is shown to be strongly influenced by plant species composition, mediated, in this case, by the provision of extra-floral nectar by one of the plant species.     

Effects of supplemental food on intra-and inter-specific behaviour of the Varied Tit Parus varius

Varied Tits Parus varius lived in flocks containing a pair, or a pair with one or two unpaired birds. To test whether the sociality of Varied Tits or their participation in mixed-species flocks is sensitive to additional food (sunflower seeds), we compared the following parameters between fed and unfed periods at the individual level: population size, degree of site fidelity, mono-specific flock size and the stability of its membership, home-range size and distribution, attendance rate with mixed-species flocks, and flock size. Neither the size of the mono-specific flock nor its membership was affected by food supply. Whether food was added or not, individuals exhibited a strong site fidelity resulting in stable population size. Supplemental feeding had no effect on home-range size or distribution. In the presence of extra food, Varied Tits were observed in mono-specific flocks but rarely in mixed-species flocks. However, when we stopped feeding, they shifted to mixed-species flocking. After adding food, mixed-species flocks were significantly smaller than in the control samples. Varied Tits were more likely to join mixed-species flocks as temperatures dropped and wind speed increased. These results suggest that intra-specific sociality of Varied Tits is relatively insensitive to food supply, but they easily shift to mixed-species flocking in relation to food and weather conditions. We conclude that Varied Tits participate in mixed-species flocks to obtain short-term benefits e.g. increased foraging efficiency but they also obtain long-term benefits from stability of pair bonds and strong site fidelity, which did not respond to supplemental food.     

One Norwegian territory of the Marsh Tit Parus palustris during 35 years

One Norwegian territory was occupied by successive pairs of Marsh Tits Parus palustris during 35 breeding seasons. Only ten males and seven females were involved. Each pair showed strict site fidelity and kept to the territory all year round. The pair bond lasted throughout life, and the duration varied from one to eight successive years. The number of successive mates varied for the males from one to two females and for the females from one to four males. Assuming that the birds were juveniles or 1 year old when ringed, they survived, on the average, 4.4 years. The oldest male survived for at least 10 years 4 months and the oldest female for 8 years 4 months. The mean survival rate of adults from one breeding season to the next was 75.8%, which is remarkably high when compared with the genera! findings in tits.     

Effects of habitat fragmentation on the timing of Crested Tit Parus cristatus natal dispersal

Crested Tit Parus cristatus young from first broods dispersed 1 week later if they were born in small isolated pine plots ('habitat fragments') compared with individuals in a large pine forest ('continuous habitat'). This delay in dispersal was caused by an extended period between fledging and dispersal. In second broods, the delay was even longer due to the interbrood interval being 9 days longer in habitat fragments. As nestlings in habitat fragments had a lower body-mass, and age at dispersal was negatively correlated with nestling body-mass within each nest, the postponed dispersal from fragments might be explained partly by a lower body-mass. Alternatively, postponed dispersal from fragments could result from a barrier effect caused by reluctance to cross inhospitable habitat.
Immigration by young from first broods into habitat fragments was delayed by approximately 3 weeks, and proportionally more second brood emigrants were recovered in this type of habitat. These results are in agreement with the hypothesis that fragments are second-choice habitat. Early immigrants into continuous habitat had a higher probability of settlement in winter flocks compared with late ones, independent of condition or age. Therefore, Crested Tits born in habitat fragments probably have a lower chance of settling in first-choice habitat.     

A model approach to project the start of egg laying of Great Tit (Parus major L.) in response to climate change

The aim of this study was to select a phenological model that is able to calculate the beginning of egg laying of Great Tit (Parus major) for both current and future climate conditions. Four models (M1–M4) were optimised on long-term phenological observations from the Ecological Research Centre Schlüchtern (Hessen/Germany). Model M1 was a common thermal time model that accumulates growing degree days (GDD) on an optimised starting date t ₁. Since egg laying of Great Tit is influenced not only by air temperature but also by photoperiod, model M1 was extended by a daylength term to give M2. The other two models, M3 and M4, correspond to M1 and M2, but t ₁ was intentionally set to 1 January, in order to consider already rising temperatures at the beginning of the year. A comparison of the four models led to following results: model M1 had a relatively high root mean square error at verification (RMSE_ver) of more than 4 days and can be used only to calculate the start of egg laying for current climate conditions because of the relatively late starting date for GDD calculation. The model failed completely if the starting date was set to 1 January (M3). Consideration of a daylength term in models M2 and M4 improved the performance of both models strongly (RMSE_ver of only 3 days or less), increased the credibility of parameter estimation, and was a precondition to calculate reliable projections in the timing of egg laying in birds for the future. These results confirm that the start of egg laying of Great Tit is influenced not only by air temperature, but also by photoperiod. Although models M2 and M4 both provide comparably good results for current climate conditions, we recommend model M4–with a starting date of temperature accumulation on 1 January–for calculating possible future shifts in the commencement of egg laying. Our regional projections in the start of egg laying, based on five regional climate models (RCMs: REMO-UBA, ECHAM5-CLM, HadCM3-CLM, WETTREG-0, WETTREG-1, GHG emission scenario A1B), indicate that in the near future (2011–2040) no significant change will take place. However, in the mid- (2041–2070) and long-term (2071–2100) range the beginning of egg laying could be advanced significantly by up to 11 days on average of all five RCMs. This result corresponds to the already observed shift in the timing of egg laying by about 1 week, due mainly to an abrupt increase in air temperature at the end of the 1980s by 1.2 K between April and May. The use of five regional climate scenarios additionally allowed to estimate uncertainties among the RCMs.