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.     

Source-sink populations in Mediterranean Blue tits: evidence using single-locus minisatellite probes

Long term studies on population biology of Blue tits (Parus caeruleus L.) in Mediterranean habitats have shown that in patchy landscapes life-history traits seem to be adapted to the predominant type of habitat, where reproductive success is higher. The “source-sink hypothesis” suggests that differences in the local production of fledglings result in an asymmetrical gene flow from rich deciduous habitats (“source”) to evergreen poor habitats (“sink”), preventing local adaptation in evergreen habitats. In this study we used single-locus minisatellite DNA probes to test the following predictions of the source-sink hypothesis: 1) source and sink populations are not genetically differentiated; 2) amount of gene flow is ranked in the following decreasing order: between source and sink habitats, among source habitats and among sink habitats; and 3) linkage disequilibrium is higher in sink than in source populations. Results were consistent with these three predictions, and with previous results obtained using other approaches. Results clearly support a source-sink functioning of Blue tit populations in southern France mosaic landscapes, and emphasise the need of combining genetic and ecological studies to understand the functioning of natural populations.     

Blue tits use selected plants and olfaction to maintain an aromatic environment for nestlings

Animals use the chemical compounds of plants as a defence mechanism against enemies, and sometimes use olfaction to discriminate and select the chemical plant substances. Some birds bring to the nest plant material that has volatile compounds that protect the host and their offspring against parasitic organisms. Here we show that blue tits on the island of Corsica ( Parus caeruleus ogliastrae ) adorn their nests with fragments of aromatic plants. These plants have chemical compounds that are used by humans to make aromatic house cleaners and herbal medicines. We also show that individual blue tits maintain an aromatic nest environment when offspring are raised, using odour cues to determine the frequency with which they replenish the nest with fresh plant material. We provide an exceptional example of the ecologically relevant use of olfaction by birds under natural conditions. To our knowledge, we present the first experimental demonstration that a free-ranging animal makes use of smell to maintain an aromatic environment for offspring with plants, supporting predictions of the nest protection hypothesis.     

Seasonal changes in a ultraviolet structural colour signal in blue tits, Parus caeruleus

Recent studies of blue tits, Parus caeruleus , have found sexual selection and a viability-indicating function of the structural ultraviolet and blue crown plumage, but the reasons for this signal variation are not understood. Furthermore, studies in England and Sweden have yielded somewhat different results (particularly with regard to the spectral position of the reflectance peak). Here we investigate whether the blue tit UV/blue ornament varies with time of year since such variation might be relevant to the signalling function as well as the apparent difference between populations. From 400 blue tits captured at two different localities in Sweden, we found that objective measures of 'hue' (spectral location), 'chroma' (spectral purity) and 'brightness' (spectral intensity), varied substantially with season. Just after moult (October), crown 'hue' is maximally UV-shifted (359 nm for males and 373 nm for females). Thereafter the peak drifts upwards and by the time of nestling feeding (June) male reflectance peaks at 404 nm and female at 413 nm. This change is probably due to feather wear as well as fat and dirt accumulation, which might constitute an additional male quality cue. Our results suggest that it is important to consider plumage age when exploring variation in structural plumage coloration, and that it can largely explain the difference between the British and Swedish studies. © 2002 The Linnean Society of London, Biological Journal of the Linnean Society , 2002, 76 , 237–245.     

Some aspects of the morphology of Blue tits (Parus caemleus) and Coal tits (Parus ater) in relation to their behaviour

An attempt is made to find functional relationships between the morphology of titmice and their behaviour. Coal tits have relatively longer feet than Blue tits, and they also have relatively long back claws and toes. This may be related to life in pine needles, the long Coal tit feet having an effect analogous to snow shoes. Blue tits, in contrast, have short strong toes which are opposable, and this may enable them to hang on oak leaves. There are small but consistent differences in leg length and proportions between Blue, Coal, Marsh and Great tits. These differences do not seem to be related to behaviour in the same way as in any other bird group. Coal tits have wings of higher aspect ratio than Blue tits, and they also have a lower wing loading. This may be related to the superior hawking and hovering ability of Coal tits. Coal tits have longer and finer bills than Blue tits, and this may be associated with small prey size and probing into crevices for food. A fine bill would be of advantage when dealing with small prey, but a long bill would be a handicap for seeing small prey close to the bill tip.     

Habitat effects on physiological stress response in nestling blue tits are mediated through parasitism

Abstract We investigated determinants of the physiological stress response mediated by stress proteins in blue tit Cyanistes caeruleus nestlings growing up in oak forests in central Spain, resulting from different forest management practices. We assessed circulating levels of the heat-shock protein HSP60 as an integrated physiological measure of the conditions experienced by nestlings during postnatal development. The effects of habitat quality and parasite infections on nestling rearing environment were then assessed through this measurement of stress response. Our results showed that newly acquired ecto- and hemoparasite infections were associated with forest habitat structural characteristics, higher prevalence of fleas and blood parasites in more mature forests, and higher prevalence of blowflies in degraded forests. While habitat characteristics did not explain variation in stress protein levels, infestation by blowfly larvae of the genus Protocalliphora and hematozoa infection by Leucocytozoon were significantly associated with higher levels of HSP60. Thus, upregulation of the expression of certain stress proteins seems to be a common physiological mechanism to alleviate the negative impact of parasite infections in growing birds. Habitat characteristics may thus indirectly determine growth conditions for forest birds mediated through their association with one of the most important selection pressures for offspring development, parasite infections.     

Stimuli for nestling begging in blue tits Cyanistes caeruleus: hungry nestlings are less discriminating

In altricial birds, nestlings usually respond to the sound and appearance of the provisioning adults by begging for food when the adults arrive at the nest. Nestlings can, however, also beg incorrectly on hearing misleading sounds in the environment and fail to beg when the adult arrives. This study uses the blue tit Cyanistes caeruleus to test the hypotheses that nestling begging strategies are influenced by the reliability of the stimulus to beg, and that nestling motivational state affects the response to different stimuli. Here, we show experimentally that nestling hunger strongly influences the response to stimuli that vary in their reliability. While hunger increases begging rate, it also increases the likelihood that nestlings will beg when the parent is absent. This is in agreement with both the predictions of signal detection theory and recent empirical work on other species. We found, however, no evidence that age-related perceptual constraints influence the begging response of ten day old nestlings to different stimuli.     

Blue tits (Cyanistes caeruleus) respond to an experimental change in the aromatic plant odour composition of their nest

Although the use of olfaction by birds is now widely recognised, the olfactory abilities of passerine birds remain poorly explored, for historical reasons. Several studies however suggest that passerines can perceive volatile compounds in several biologically relevant contexts. In Corsica, recent findings suggest that cavity-nesting blue tits may use volatile compounds in the context of nest building and maintenance. Although they build their nests mainly from moss, female blue tits also frequently incorporate fragments of several species of aromatic plants in the nest cup. In field experiments, breeding female blue tits altered their nest maintenance behaviour in response to experimental addition of aromatic plants in their nest. In aviary experiments, captive male blue tits could be trained to detect lavender odour from a distance. Here I report results from a field study aimed to test whether adult blue tits altered their chick-feeding behaviour after an experimental change in nest odour composition. I experimentally added fragments of aromatic plant species that differed from those brought in the nests before the start of the experiment in a set of experimental nests and added moss, the basic nest material, in a set of control nests. Both male and female blue tits hesitated significantly longer entering the nest cavity after addition of new aromatic plant fragments, as compared to moss addition. This response was especially observed during the first visit following the experimental change in nest plant composition. Nest composition treatment had no effect on the time spent in the nest. This study demonstrates that free-ranging blue tits detect changes in nest odour from outside the nest cavity.     

Food storing by marsh tits

Wild marsh tits (Parus palustris) were allowed to hoard radioactively labelled sunflower seeds, which were subsequently found using a portable scintillation counter. Seeds were stored singly, in various sites close to the ground. Different birds favoured different types of site, although this preference was changeable. Seed density decreased with increasing distance from the feeder, and there was a negative correlation between seed sequence number and the distance it was carried. Hoarded seeds disappeared more rapidly than control seeds in identical sites 100 cm away, suggesting that the birds remember their exact location. Seeds also tended to be stored and recovered in the same sequence. The adaptive significance of these results is discussed.     

Snake-like calls in breeding tits

Hole-nesting tits belonging to the family Paridae produce a hissing display that resembles the exhalatory hiss of a snake. When a predatory animal enters the nest hole of a tit, tits often hiss vigorously, while lunging their head forward and shaking their wings and tail, until the intruder retreats. We assessed the acoustic similarity between such hiss calls from 6 species of tits, snake hisses, and tit syllables used in alarm vocalizations, as well as white noise as a control. Tit hiss calls showed a high degree of similarity with snake hisses from 3 different snake families. Tit hisses had lower similarity to syllable alarm calls, suggesting convergence of tit hisses in their spectral structure. Hiss calls would only be effective in protecting nest boxes if nest predators responded to these calls. In order to test this hypothesis, we trained individual Swinhoe’s striped squirrels, Tamiops swinhoei hainanus, a common predator of egg and nestling tits, to feed at feeders in proximity to nest boxes. We compared the aversive response of squirrels to tit’s hiss calls and white noise, presented in random order. Squirrels showed a higher degree of avoidance of feeders when hiss calls were played back than when white noise was presented. In conclusion, our study suggests that hole-nesting birds have evolved convergent snake-like hiss calls, and that predators avoid to prey on the contents of nest boxes from which snake-like hisses emerge.     

Night-time behaviour of egg-laying tits

We provide evidence that nestbox-breeding tits roost in their boxes at night during the egg-laying period, but are able to do so while avoiding initiating incubation, despite choosing to settle into the nest-cup. Nest-cup temperatures of Great Tits Parus major , during the egg-laying period, were usually below the threshold for embryo development. This suggests an ability to control heat transfer to the eggs in spite of the close proximity of the female to her clutch. We suggest that an important benefit of roosting in the nest-cup during the egg-laying period is reduced thermoregulatory costs, which may be important in moderating the energetic costs of egg production. Another possible advantage is that developmental abnormalities associated with egg-chilling are avoided.     

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.     

Mutagenicity of ultraviolet A radiation in the lacI transgene in Big Blue mouse embryonic fibroblasts

Sunlight ultraviolet A (UVA) irradiation has been implicated in the etiology of human skin cancer. A genotoxic mode of action for UVA radiation has been suggested that involves photosensitization reactions giving rise to promutagenic DNA lesions. We investigated the mutagenicity of UVA in the lacI transgene in Big Blue mouse embryonic fibroblasts. UVA irradiation of these cells at a physiologically relevant dose of 18J/cm(2) caused a 2.8-fold increase in the lacI mutant frequency relative to control, i.e., 12.12+/-1.84 versus 4.39+/-1.99 x 10(-5) (mean+/-S.D.). DNA sequencing analysis showed that of 100 UVA-induced mutant plaques and 54 spontaneously arisen control plaques, 97 and 51, respectively, contained a minimum of one mutation along the lacI transgene. The vast majority of both induced- and spontaneous mutations were single base substitutions, although less frequently, there were also single and multiple base deletions and insertions, and tandem base substitutions. Detailed mutation spectrometry analysis revealed that G:C-->T:A transversions, the signature mutations of oxidative DNA damage, were significantly induced by UVA irradiation (P<0.003). The absolute frequency of this type of mutations was 7.4-fold increased consequent to UVA irradiation as compared to control (3.38 versus 0.454 x 10(-5); P<0.00001). These findings are in complete agreement with those previously observed in the cII transgene of the same model system, and reaffirm the notion that intracellular photosensitization reactions causing promutagenic oxidative DNA damage are involved in UVA genotoxicity.     

Levels of extra‐pair paternity are associated with parental care in penduline tits (Remizidae)

In most passerine birds, individuals attempt to maximize their fitness by providing parental care while also mating outside their pair bond. A sex‐specific trade‐off between these two behaviours is predicted to occur, as the fitness benefits of extra‐pair mating differ between the sexes. We use nest observations and parentage analysis to reveal a negative association between male care and the incidence of extra‐pair paternity across three species of penduline tit (Remizidae). This provides evidence of a trade‐off between these two behaviours, possibly due to the devaluing of paternal care by extra‐pair offspring.     

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.     

Behavioral snake mimicry in breeding tits

Many animals mimic the behavior or the appearance of venomous snakes.When humans or other potential predators place their hand near the nest of tits belonging to the family Paridae(and a few other species),the incubating female performs a hissing display that mimics the inhalation hiss of a viper or another snake.They hiss vigorously while lunging their head forward and shaking their wings and tail,repeating this behavior several times.The structure of the hiss in tits is similar to that of the inhalation hiss of a snake,providing evidence of significant convergence of the mimic toward the model.The behavior of individual females is repeatable among tri als.Individuals that flew away from their nest box only performed the hissing display on 6%of later trials,when present at their box,whereas individuals that did not fly away hissed on 28% of occasions,consistent with great tits Parus major either cautiously flying away or staying put on their nest while actively defending it.Individuals that flew away produced fewer chicks than individuals that stayed and hissed.The hissing display was more common when snakes were more abundant:1)When breeding late during the season;2)when breeding at sites with more snakes;and 3)when breeding in subtropical and tropical China with a higher abundance of snakes than in Denmark with a lower abundance.The frequency of nest predation was higher in sites with no snakes,and the frequency of predation increased with decreasing frequency of hissing display.These findings are consistent with expectations for frequency-dependent selection acting on snake mimicry.