Prey Detection in Two Tit Species,Parus ater and P. cristatus

We studied the prey-detection response of two related tit species (Parus ater, coal tit, and P. cristatus, crested tit) which occur in similar habitats and which are separated by microhabitat use and mode of locomotion. Specifically, we tested whether the two species differ in their ability to detect cryptic prey, with the hypothesis that coal tits perform better because of their longer stays in microhabitat patches. We expected that interspecific differences and the flexible responses within species to different levels of prey crypticity both depend on search intensity. In the experiments, tits had to forage in four different prey conditions: conspicuous + small, conspicuous + large, cryptic + small, and cryptic + large. Capture success, the time needed to detect prey, and stare duration were measured. Both species were less successful with cryptic than with conspicuous prey and did worst with small and cryptic prey. They also needed more time for detection and stared longer when prey was cryptic. When prey had been overlooked, preceding stare durations were shorter than those ending in successful searches. There were clear differences between the species in performance, with the crested tit doing worse. Coal tits, on average, were more successful in detecting prey, returned less frequently to already emptied patches, detected prey faster and yet they stared more briefly. The within-species results support current notions on predator search, but the differences between species cannot be explained with the same mechanisms. We discuss these findings in the context of the foraging ecology of these species and conclude that crested tits use different cues to detect prey and/or search a larger area per unit time.     

Hoarding of individual nuthatches Sitta europaea and march tits Parus palustris

The hoarding behaviour of one pair of nuthatches Sitta europaea L. and of four individual marsh tits Parus palustris L. was studied during autumn and winter in a deciduous grove in central Sweden. Sunflower seeds were offered at a feeding table. AM individuals studied stored seeds intensively throughout the whole study period. The two species showed very different hoarding patterns. The nuthatch pair aggressively defended the area around the feeding table, which was within their territory, from other nuthatches and also seemed to affect the hoarding behaviour of the coexisting marsh tits. Some significant differences were found also between con-specifics in some aspects of hoarding behaviour, which could in some cases be related to sex or dominance. These differences may make hoarding profitable for the individual birds if they also differ in their foraging patterns individually, and so will have a greater probability of finding their own stored items than other individuals. However, the evidence for individual segregation of hoarding patterns is not conclusive.     

Great tits (Parus major) searching for artificial prey: implications for cryptic coloration and symmetry

It has been suggested that bilateral symmetry may impose a costfor animals relying on camouflage because symmetric color patternsmight increase the risk of detection. We tested the effect ofsymmetry on crypsis, carrying out a predation experiment withgreat tits (Parus major) and black-and-white–patterned,artificial prey items and background. First, we found that detectiontime was significantly longer for a highly cryptic, asymmetricpattern based on a random sample of the background than forits symmetric variants. Second, we were able to arrange theelements of a prey pattern in a way that the resulting asymmetricpattern was highly cryptic and, furthermore, its symmetric variantwas highly cryptic as well. We conclude that symmetry may imposea substantial cost on cryptic patterns, but this cost variesamong patterns. This suggests that for prey, which predatorstypically view from an angle exposing their symmetry, selectionfor pattern asymmetry may be less important and selection fordecreased detectability cost of symmetry may be more importantthan previously thought. This may help to understand the existenceof so many prey with cryptic, symmetric color patterns.     

Ecological and morphological variation of Canary Island blue tits, Parus caeruleus (Aves: Paridae)

Canary Island blue tits have differentiated within the last million years. Compared with mainland populations they have short wings, large beaks and tarsi, and tend to be less variable in these parameters. Within the archipelago the eastern populations have the shortest wings and tarsi but the widest and deepest beaks. The eastern populations may have been derived from central island ones, following extinction of the original eastern populations during a warm period in the last Ice Age. New measurements have been used to test contrasting predictions of hypotheses that explain inter-island morphological variation in terms of either climatic or ecological influences. Neither hypothesis is sufficient to explain the variation. A model which combines mem is proposed as follows. Mild winter temperatures have selected for small body size and wing length, especially on the hot, dry, eastern islands. On the other pine-forested, islands, selection has favoured relatively long tarsi and long, slender, beaks for efficient foraging among pine foliage. The absence of die pine-dwelling coal tit from the islands has facilitated mis evolutionary shift. It is shown that blue tits in pines forage more in the needles and cones on the Canary Islands than they do on die North African mainland.     

Predator-induced take-off strategy in great tits (Parus major)

When birds are attacked by predators the initial take-off is crucial for survival. The strategy in the initial phase of predator evasion is probably affected by factors such as body mass and presence of cover and conspecifics, but it may also be a response to the character of the predator''s attack. In choosing an angle of flight, birds face a trade-off between climbing from the ground and accelerating across the ground. This is, to our knowledge, the first study investigating whether the attack trajectory of a raptor affects the take-off strategy of the prey bird. First-year male great tits (Parus major) adjusted take-off angle to a model predator''s angle of attack. Birds attacked from a steep angle took off at a lower angle than birds attacked from a low angle. We also compared take-offs at dawn and dusk but could not find any measurable effect of the diurnal body mass gain (on average 7.9%) in the great tits on either flight velocity or angle of ascent.     

Behavioral responses to conspecific mobbing calls are predator‐specific in great tits (Parus major)

When facing a predator, animals need to perform an appropriate antipredator behavior such as escaping or mobbing to prevent predation. Many bird species exhibit distinct mobbing behaviors and vocalizations once a predator has been detected. In some species, mobbing calls transmit information about predator type, size, and threat, which can be assessed by conspecifics. We recently found that great tits (Parus major) produce longer D calls with more elements and longer intervals between elements when confronted with a sparrowhawk, a high‐threat predator, in comparison to calls produced in front of a less‐threatening tawny owl. In the present study, we conducted a playback experiment to investigate if these differences in mobbing calls elicit different behavioral responses in adult great tits. We found tits to have a longer latency time and to keep a greater distance to the speaker when sparrowhawk mobbing calls were broadcast. This suggests that tits are capable of decoding information about predator threat in conspecific mobbing calls. We further found a tendency for males to approach faster and closer than females, which indicates that males are willing to take higher risks in a mobbing context than females.     

The influence of brood size upon metabolic rate and body temperature in nestling Blue tits Parus caeruleus and House sparrows Passer domesticus

Metabolic rate and body temperature in nestling Blue tits and House sparrows were measured in broods of different size and age. Surface-volume ratio effects were found in both poikilothermic and homoiothermic Blue tits at ambients of 15°C but not at 20°C. The possibility of incipient hyperthermia amongst young nestlings maintaining sub-adult body temperatures is discussed. For the House sparrows heat retention by the nest was of greater importance than the surface-volume effect. Differences in nest structure between the two species are described and related to the thermal requirements of nestlings in large as against small broods.     

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.     

Hissing calls improve survival in incubating female great tits (Parus major)

Nest predation is among the most important selective pressure shaping nest-site selection and nest defense behavior in many avian species. In this study, we tested whether the production of one such nest defense behavior—hissing calls—may improve survival of incubating female great tits (Parus major). We found that 72.5 % of incubating females gave hissing calls when they were exposed to a stuffed woodpecker in their nest boxes. The repeatability of the number of hissing calls given was high, as was the latency to give the call. Additionally, natural nest predators attacked hissing and nonhissing females equally often. However, hissing females survived better than silent females. We tested responses of feral cats to playbacks of hissing call during their attacks of nest boxes and found that hissing calls prevented the predator attacks. Taken together, our findings indicate that hissing calls can deter predator attacks and potentially increase survival rates of nesting great tits or their offspring, or both. The propensity to give hissing calls may be related to personality type of incubating female great tits, which needs to be tested experimentally.     

Responses of great tits Parus major to small tortoiseshells Aglais urticae in feeding trials; evidence of aposematism

Abstract.  1. The small tortoiseshell Aglais urticae (Lepidoptera, Nymphalidae) is one of several red-coloured butterflies and moths that are active in early spring in Norway. It has been suggested that tortoiseshells may be warningly coloured and unpalatable to birds, however no experiments have been carried out to test this hypothesis.
2. The reactions of wild-caught great tits Parus major (Paridae) when offered dead specimens of the tortoiseshell and four palatable control species were studied. Two experiments were carried out. In expt 1, only intact prey was presented to the birds. In expt 2, the birds were offered only experimentally de-winged prey. Hence, in expt 2, it was possible to explore to what extent the birds' reaction to the prey offered to them in expt 1 could be explained as a reaction to visual stimuli from the wings of the prey.
3. In both experiments, tortoiseshells were attacked hesitantly, eaten slowly, and rejected often compared with controls, providing evidence that tortoiseshells are mildly distasteful to great tits. Moreover, all birds were more hesitant to attack intact than de-winged tortoiseshells, while there was no such effect for control prey, indicating a warning effect associated with the butterfly wings. There were also indications of avoidance learning in the birds to the presented tortoiseshells. These results are consistent with the idea of aposematism.     

Foraging behaviour of individual coal tits,Parus ater,in relation to their age,sex and morphology

The aim of this study was to reveal the causes and consequences of individual differences in foraging behaviour of coal tits, Parus ater, on the island of Gotland in the Baltic. On Gotland the willow tit, P. montanus, and the crested tit, P. cristatus, are absent, while these larger species are abundant competitors on the mainland. The Gotland population of coal tits exploited parts of the foraging niches of their absent competitors. Gotland coal tits are bigger than those on the mainland; they have thus evolved towards the size of the missing competitors. Individual differences in niche use were related to their morphology. Large birds more often used the inner parts of trees whereas small individuals foraged more on needles in the outer parts of the trees. Also, individuals with wing shapes more suitable for hovering and slow flight foraged more on needles. It is argued that morphology affects the profitability of different foraging sites. However, for foraging site selection, age was more important than morphology. Older, and probably more dominant, individuals occupied the most profitable foraging sites.     

Does foraging behaviour explain the poor breeding success of great tits Parus major in northern Europe?

Great tits Parus major have generally much poorer breeding success in northern Finland than in mid- and western Europe. The aim of this study is to find out whether the poor success is linked to foraging behaviour. This was studied by monitoring great tits' foraging behaviour and food abundance in different substrates during the breeding cycle in two populations, N Finland (65°N) and Latvia (56°N). It was shown that breeding success, from eggs to fledglings, was significantly poorer (56% vs. 93%) and caterpillar abundance considerably lower in the northern population. The general patterns in foraging behaviour were the same in N Finland and Latvia (especially the preference of birches Betula spp.) indicating that great tits used basically the same species-typical foraging strategy in both populations. However, thin branches of birch were preferred in the north but avoided in Latvia, which may suggest that northern great tits have changed their foraging niche towards the outer parts of canopy, a niche typically occupied by the blue tit. This shift is theoretically advantageous, since the outer parts of the canopy are the richest caterpillar source. In practice, however, the poorer success of great tits indicates that this is not a beneficial strategy. Primarily, great tits in the northern population seem to be food limited since they lay too large clutches in relation to accessible food resources. This may be because great tits are not adapted to lower caterpillar production in the northern margins of their distribution and cannot change their narrow diet (3/4 caterpillars), like e.g. willow tits can. In search for caterpillars, food accessibility is further limited because the great tits' normal foraging behaviour, with wide search radius, may not function properly in the denser, outer parts of the canopy. Great tits may also be too heavy to forage efficiently on leafed twigs.     

Body size, prey handling efficiency and choice of diet in three coniferous forest tits (Paridae)

The main purpose of this study was to link morphological differences between great tit ( Parus major ), willow tit ( P. montanus ) and coal tit ( P. ater ) and their rate of energy acquisition and choice of diet in order to explore the potential for competitive relations between them more directly. Handling times were measured in the laboratory by presenting mealworms of different sizes to the birds. Great tits were more efficient in handling large prey than were the smaller-bodied willow- and coal tits; for small prey sizes the coal tit was the least efficient species. Using the ratio of prey mass to the handling time value, a utility function for each species was constructed. These results suggests a potential for a segregation of the species on the food axis. However, results from the prey choice experiment show that despite considerable differences in functional morphology between the three species they do not differ significantly in the range of prey size exploited. My results suggest that the alleged importance of prey size partitioning is not likely to play the major role for the coexistence of these coniferous forests tits.     

Natal dispersal and personalities in great tits (Parus major)

Dispersal is a major determinant of the dynamics and genetic structure of populations, and its consequences depend not only on average dispersal rates and distances, but also on the characteristics of dispersing and philopatric individuals. We investigated whether natal dispersal correlated with a predisposed behavioural trait: exploratory behaviour in novel environments. Wild great tits were caught in their natural habitat, tested the following morning in the laboratory using an open field test and released at the capture site. Natal dispersal correlated positively with parental and individual exploratory behaviour, using three independent datasets. First, fast-exploring parents had offspring that dispersed furthest. Second, immigrants were faster explorers than locally born birds. Third, post-fledging movements, comprising a major proportion of the variation in natal dispersal distances, were greater for fast females than for slow females. These findings suggest that parental behaviour influenced offspring natal dispersal either via parental behaviour per se (e.g. via post-fledging care) or by affecting the phenotype of their offspring (e.g. via their genes). Because this personality trait has a genetic basis, our results imply that genotypes differ in their dispersal distances. Therefore, the described patterns have profound consequences for the genetic composition of populations.     

Behavioural responses to changing landscapes: flock structure and anti-predator strategies of tits wintering in fragmented forests

We analysed the effects of forest fragmentation on the flock structure of insectivorous forest passerines (Parus, Aegithalos, Certhia, Regulus, etc.), and on the anti‐predator behaviour and energy management of blue tits in these flocks. We surveyed flocks in Central Spain during two winters. Flocks in fragments comprised fewer individuals and species than flocks in unfragmented forests. The most abundant species in forest flocks (blue tit, Parus caeruleus, and firecrest, Regulus ignicapillus) were also the most abundant in fragments, while the rarest species in the area never occurred in small woodlots. We investigated how fragmentation and related changes in flock structure affect anti‐predator behaviour of blue tits, a widely distributed species in the area. In fragments but not in forests, blue tits increased scanning rates with decreasing flock size. Vigilance was relaxed when great tits, Parus major, were abundant as flock mates, suggesting that the absence of this dominant species in fragments could intensify anti‐predator behaviour of blue tits. Blue tits enhanced anti‐predator behaviour in the second winter parallel to an increase in the abundance of raptors. This behavioural change was stronger in fragments, where blue tits foraged deeper in the canopy and increased scanning and hopping rates. Under increased predation risk, birds are expected to reduce body mass to improve predator avoidance. On average, blue tits weighed similar in fragments and forests the second winter. However, they accumulated fat along the day in fragments only, and adjusted body mass to body size more closely in that habitat type. This suggests that blue tits perceived fragments as unpredictable habitats where fattening would help avoid starvation, but also as dangerous sites where overweight would further increase the risk of predation. In summary, our results support that fragmentation affects individual behaviour of blue tits, and show the potential of behavioural approaches to unravel how different species face the advancing fragmentation of their habitats.     

Interspecific aggression declines seasonally in breeding great tits Parus major

During the breeding season, great tits show aggression to protect their nest from intra‐ and interspecific intruders. Aggression is a labile trait that can be plastically expressed as a result of individual differences (e.g., personality), seasonal gradients in the costs and benefits of aggression, or other environmental components (e.g., number of competitors). Competitors may try to take over great tit nests, because the number of suitable nesting sites is limited, and great tits may guard high quality territories. Taking over a great tit nest may be especially fruitful in early phenological stages (egg laying) when great tits frequent their nests less often. However, great tits may compensate for this vulnerability by being more aggressive toward intruders during early nesting stages, a pattern that has already been established in an intraspecific context. Previous studies have shown that interspecific intruders were most likely to die from great tit aggression during great tit egg laying, suggesting great tits may also be more aggressive during this phase in an interspecific context. Here, I tested this hypothesis with simulated territorial intrusions in great tit territories using taxidermized blue tits Cyanistes caeruleus (hereafter called blue tit models). Great tit aggression (number of calls and approach distance toward blue tit model) was assayed during egg laying, incubation, and chick rearing in the breeding season of 2014. Although sample size was low due to a high fraction of non‐responders (n = 44 out of 89 assays across 26 out of 35 individuals), I found that great tits showed a seasonal decline in aggressiveness, which is congruent with intraspecific results on this study species. I discuss my findings in the context of differential adjustment to climate change between interspecific competitors.     

Personality in Blue Tits (Cyanistes caeruleus) and its Effect on Their Breeding Success

In this study, we investigated whether blue tits (Cyanistes caeruleus) showed consistent variation in neophobia responses. The experiment represents a test of how personality, consistent individual differences in behaviour, may have fitness consequences. The experimental design consisted of a control observation followed by two neophobia tests, during which the parent's behaviour was recorded using the instantaneous sampling method. Blue tits showed consistent variation in neophobia responses, which was related to total offspring biomass, but not to mean offspring weight or fledging success. General linear mixed models analyses suggest that there may be ideal neophobia trait combinations in breeding pairs that maximise breeding success.     

Influence of radar radiation on breeding biology of tits (Parus sp.)

The aim of the present study is to observe the influence of long-term exposure to radar radiation on breeding biology of tits (Parus sp.), living and building nests around a military radar station, emitting pulse-modulated microwave radiation of 1,200-3,000 MHz. Two series of 36 nest-boxes each were located on the radar station area. Measurements of exposure were performed separately for each nest-box. Average power density (P(av), W/m(2)) and dose of exposure (W/m(2) x h) were recorded for each nest-box during 45 days. Control nest-boxes (N = 42) were located in other part of the same forests, free from radar radiation. The assessment of effects of radar exposure on breeding biology of tits included number of inhabited nest-boxes, number of eggs, and nestlings in the nest (Why not chick mortality?). Experimental nest-boxes were either exposed to relatively high levels of radiation (2.0-5.0 W/m(2), mean 3.41 +/- 1.38 W/m(2)) or an intermediate level of radiation that ranged from 0.1-2.0 W/m(2) (mean 1.12 +/- 0.84 W/m(2)). For control nest-boxes the exposure ranged from 0.001-0.01 W/m(2) (mean 0.0062 +/- 0.0007 W/m(2)). Only blue or great tits occupied all nest-boxes, used in the experiment. The number of nesting blue tits was higher in nest-boxes located on the radar station area than in the control boxes. In contrast, control nest-boxes were inhabited mainly by great tits. On the radar station area, blue tits nested in high exposed nest-boxes (67,0%) and great tit occupied mainly these boxes, which were exposed to low-level radiation (62,5%), the difference being statistically significant (p < 0.01). No statistically significant differences in other parameters of breeding biology (number of eggs per box, number of nestling per box) were observed between tits occupying exposed and control nest boxes. Results of the present study show that radar radiation generally does not lead to decrease of number of nesting tits, but may cause shifts in tits species living around the radar station. (But is the microhabitat, apart from the radiation level, around each nest box more likely to attract one species of tit or another?).     

Hissing females of great tits (Parus major) have lower breeding success than non‐hissing individuals

Antipredator strategies vary remarkably between individuals within populations. Parents tend to take greater risks when brood value is increased. Moreover, individuals consistently differ in a whole suite of correlated behaviours that may cause distinctive responses to predators. It is likely that individual differences in antipredator behaviour may co‐vary with proxies for fitness such as reproductive success. We used a 4‐year data from wild great tits (Parus major) to test whether passive and active antipredator strategies (females with no response vs. those giving hissing calls towards a nest predator) during the incubation stage can reflect variation in breeding success. Although clutch size did not depend on hissing behaviour, the number of surviving offspring from eggs and neonates to fledglings was higher for non‐hissing than hissing birds. We conclude that females with distinct antipredator strategies can prioritize different fitness components.