Septum volume and food‐storing behavior are related in parids

The hippocampal formation (HF) of food‐storing birds is larger than non‐storing species, and the size of the HF in food‐storing Black‐Capped Chickadees (Poecile atricapillus) varies seasonally. We examined whether the volume of the septum, a medial forebrain structure that shares reciprocal connections with the HF, demonstrates the same species and seasonal variation as has been shown in the HF. We compared septum volume in three parid species; non‐storing Blue Tits (Parus caeruleus) and Great Tits (Parus major), and food‐storing Black‐Capped Chickadees. We found the relative septum volume to be larger in chickadees than in the non‐storing species. We also compared septum and nucleus of the diagonal band (NDB) volume of Black‐Capped Chickadees at different times of the year. We found that the relative septum volume varies seasonally in food‐storing birds. The volume of the NDB does not vary seasonally. Due to the observed species and seasonal variation, the septum, like the hippocampal formation of food‐storing birds, may be specialized for some aspects of food‐storing and spatial memory. © 2002 Wiley Periodicals, Inc. J Neurobiol 51: 215–222, 2002     

Yellow plumage colour of Great Tits Parus major correlates with changing temperature and precipitation

Weather is known to affect the phenology and behaviour of birds, but weather-related changes of phenotypic traits involved in communication have received little attention. Using an 8-year dataset, we investigated links between carotenoid-based reflectance of the freshly moulted breast of Great Tits Parus major, weather during the moulting period and food availability during the preceding breeding season, and we investigated interannual changes. In both sexes, we found a change of colour expression to more saturated and darker yellow over the study period in parallel with increasingly dry and warm weather during moult. These results indicate that the expression of traits playing roles in communication may be controlled by weather and may therefore shift in response to changing climate.     

The effect of dominance rank on fat deposition and food hoarding in the Willow Tit Parus montanus– an experimental test

We studied the effects of dominance rank on fat deposition and hoarding behaviour in Willow Tits Parus montanus . Dominant individuals can displace subordinates which gives them priority to new food sources; they can also pilfer stored food from subordinates. This gives subordinates less certain access than dominants both to their own caches and to new food sources. Theory predicts that subordinates should invest more than dominants both in body fat reserves and stored food. Empirical evidence is equivocal; some studies have shown that subordinates built up larger reserves than dominants, whereas others show the opposite. In an earlier indoor experiment, Pravosudov and Lucas found no effect of rank on either hoarding rate or fat reserves, but the experimental design was such that the results were ambiguous. This paper reports on a similar, but improved, experiment in outdoor aviaries. However, our results agree with the earlier experiment, since we found no effect of rank on either food storing or fat deposition. The reasons for this are explored.     

Coal Tit fat-storing patterns during the non-breeding season: the role of residence status

According to the optimal body mass hypothesis, resident individuals, having priority of resource access and better knowledge of food availability in an area, should carry smaller fat reserves than transients whose resource predictability is lower. We tested this prediction in a free-living population of Coal Tits Parus ater in sub-alpine coniferous forest during three winters. We examined the role of residence status in determining the daily patterns of fat accumulation of individuals using time of day, temperature, season, year and body size as covariates. Only time of day, residence status and temperature significantly affected daily variation of fat score in Coal Tits. Fat scores were higher in transients than residents, although within residents, juveniles did not carry significantly more fat than adults. Our results show that fattening strategies are associated with residence status in the Coal Tit, in support of the hypothesis that transient individuals facing lower food predictability carry larger body fat reserves. The finding that within residents, patterns of fat accumulation did not differ between adults and juveniles, suggests that residence per se , involving higher food predictability, rather than age-related dominance or hoarding behaviour, determines fattening strategies in wild Coal Tits.     

Food resource matching by foraging tits Parus spp. during spring-summer in a Mediterranean mixed forest; evidence for an ideal free distribution

We analysed whether patterns of microhabitat use by Blue Tits Parus caeruleus , Great Tits Parus major and Crested Tits Parus cristatus inhabiting a mixed forest consistently matched the patterns of food availability experienced by foraging birds during spring-summer. The use of five microhabitats by each bird species (the foliage of three tree species, shrubs and ground) and the availability of food in trees during the prebreeding, breeding and post-breeding periods of the birds' annual cycle were measured. All three tit species foraged mainly in the outer part of tree canopies (small branches and leaves or needles). Tit distributions between tree species matched food resource distributions irrespective of overall food resource levels, which varied four-fold between the study periods, and tit species. Tits also exploited secondary microhabitats (shrubs and ground) in periods of low food availability; Blue Tits tended to use shrubs, whereas Great and Crested Tits foraged on the ground. Between-trees distributions fitted that expected from an ideal free distribution, suggesting that food availability and intraspecific exploitative competition were the main factors governing tree use by tits. In contrast, patterns of use of secondary microhabitats (shrubs and ground) seemed to indicate a role for the species-specific morphological configurations of each tit species since Blue Tits are better adapted to hang and tended to forage in shubs, whereas Great and Crested Tits are better adapted to feed on horizontal surfaces and tended to forage on the ground. No evidence of interspecific interactions was observed. Overall, the results pointed to an independent exploitation of Mediterranean mixed forest by each bird species, food availability and food accessibility being the main factors affecting microhabitat use by foraging tits.     

Foraging behaviour of Blue Tits Parus caeruleus in a patchy environment under contrasting levels of natural food supply

We analysed the foraging behaviour of free-ranging Blue Tits Parus caeruleus in open holm oak Quercus ilex woodlands of western Spain during winter. Such woodlands are patchy for foraging tits because of the scattered distribution of trees and the patterns of abundance of canopy arthropods within and among trees. Results were compared with those obtained in spring of the same year, when we found that the foraging behaviour and spatial distribution of Blue Tits were largely unaffected by food availability (Pulido and Díaz 1997). Patch (tree) residence time was highly variable both within and among individual birds, and it was uncorrelated with either previous travel time or patch quality. Contrary to a priori expectations, the behaviour of tits did not conform to short-term energy maximizing rules in winter, in spite of a 2.5-fold decrease in food supply from spring to winter and a likely 2-fold increase in bird requirements. Instead, birds tended to fly towards patches that were further away than locally available. Overall, we conclude that energy intake rate was not the fitness-related currency that birds were trying to maximize while foraging.     

Nectar-feeding by suburban Blue Tits: contribution to the diet in spring

The incidence of nectar feeding by 2 pairs of Blue Tits Parus caeruleus was investigated in relation to the availability of nectar and alternative food. Nectar was not the most preferred food and the occurrence of nectar feeding did not correlate significantly with most measures of nectar availability, but the Blue Tits selected the most productive flowers. Nectar feeding frequently occurred when the preferred peanuts were unavailable due to interspecific competition. The nectar resource was under-utilized by the Blue Tits except at the beginning of the flowering period. Despite this, nectar was a highly profitable food source, yielding 0.33-0.38 kJ min^?1 foraging. During the flowering period nectar was estimated to contribute up to 32.7% of the average daily metabolic rate (ADMR) of the male and up to 49.3% of the ADMR of the female, with means over the 2 years of 7.4% and 13.2% per day for the male and female, respectively. The importance of nectar in the ecology of Blue Tits is discussed.     

The ecology of the avian brain: food-storing memory and the hippocampus

Some species of birds store food, often hoarding several hundreds of seeds over a period of just a few weeks. Field and laboratory studies have demonstrated that food-storing species have an impressive memory and an enlarged region of the brain, the hippocampal region. Lesion experiments have shown that the hippocampus is important in accurate retrieval of stored food. Taken together, these results have led to the hypothesis that the enlarged hippocampus is associated with the memory requirements of retrieving stored food. In this review, we discuss four areas of study: comparative studies of the brain, comparative studies of behaviour, developmental plasticity and seasonal changes in food storing and the hippocampus.     

The ecology of the avian brain: food-storing memory and the hippocampus

Some species of birds store food, often hoarding several hundreds of seeds over a period of just a few weeks. Field and laboratory studies have demonstrated that food-storing species have an impressive memory and an enlarged region of the brain, the hippocampal region. Lesion experiments have shown that the hippocampus is important in accurate retrieval of stored food. Taken together, these results have led to the hypothesis that the enlarged hippocampus is associated with the memory requirements of retrieving stored food. In this review, we discuss four areas of study: comparative studies of the brain, comparative studies of behaviour, developmental plasticity and seasonal changes in food storing and the hippocampus.     

Geographic variation of the storing behaviour in the Coal tit Parus ater: role of winter residency and environmental conditions

Foraging and hoarding behaviour of the Coal Tit Parus ater were studied in two widely separated populations, in the Pyrenees and in Norway. The two populations differed in their degree of winter residence and environmental pressures. Location of foraging and hoarding sites were described in the Pyrenees during winter, and during autumn in Norway where no hoarding takes place during winter. Significant differences in the hoarding behaviour of Coal Tits between these areas were observed. In the Pyrenees, birds hoarded on inner and lower parts of the trees, using trunks and thick branches as caching sites, which are the substrates most heavily used in conditions of winter environmental stress. In Norway, birds hoarded items on outer and higher parts of the trees, mainly among needled twigs, which were the substrates used in normal foraging. Animal food was hoarded more frequently by the Norwegian population. Our results show that hoarding behaviour varies among populations and suggest that the differences recorded may be related to differences in competitive pressures and the prevailing environmental conditions. Furthermore, the absence of a safe long-term hoarding niche in the Norwegian population, due to interspecific competition, would make a strong sedentary habit difficult, offering a functional explanation for the differences observed in the degree of winter residence between Norwegian and Pyrenean Coal Tits.     

THE SELECTION OF TITS PARUS SPP. BY SPARROWHAWKS ACCIPITER NISUS

Selection of the Wytham Wood population of tits by Sparrowhawks during the tit post-fledging period was studied over three years. Within three weeks of the median date of tit fledging, juvenile Blue Tits were selected more frequently than juvenile Great Tits but thereafter juvenile Great Tits were selected more frequently; it is suggested that the former was due to greater vulnerability of Blue Tits in the two to three weeks after fledging, and the latter due to changes in availability due to immigration/emigration of juvenile tits. When juvenile tits killed by hawks were compared on the basis of brood and physical characteristics with their cohorts in the entire population and in those surviving to the following year, only one factor, fledging date, was found to have affected selection by hawks. It is believed that this was because hawks selected late fledged young still in family parties in preference to early fledged young which had already become independent of their parents and were foraging in the better cover of the forest canopy. When adults killed by hawks were compared with all adults available for selection on the basis of species, sex, timing of nesting and whether or not they had been born in the wood, the only selection trend found was that male Great Tits were taken more often than females due, possibly, to greater risk of exposure for males while foraging or during territorial behaviour. When compared with availability, juveniles were selected more frequently than adults in only one year, possibly a result of yearly differences in the number of prey available per hawk, the rate of non-predator related juvenile mortality or unknown hawk hunting strategies. The overall finding of relative non-selectivity by hawks was attributed to the surprise factor associated with the hunting methods used by Sparrowhawks in woodland.     

Role of extrinsic factors in the control of clutch-size in the Blue Tit Parus caeruleus

Extrinsic factors that play a role in the control of clutch-size in Blue Tits Parus caeruleus were investigated. The disruption of ovarian follicular growth, and hence the cessation of egg-laying, is caused by a stimulus generated by eggs present in the nest. The timing of this disruption varies among females during the laying period and may happen as late as after the laying of the eighth egg of the clutch; the larger the clutch, the later the disruption of follicular growth. The timing of follicular disruption is also related to ambient temperature because the proportion of female Blue Tits that cease to lay increases when higher temperatures occur toward the end of laying. It is suggested that the use of temperature to control clutch-size may be adaptive because the food peak on which parents rely to feed their chicks is likewise affected by temperature.     

Seasonal hippocampal plasticity in food-storing birds

Both food-storing behaviour and the hippocampus change annually in food-storing birds. Food storing increases substantially in autumn and winter in chickadees and tits, jays and nutcrackers and nuthatches. The total size of the chickadee hippocampus increases in autumn and winter as does the rate of hippocampal neurogenesis. The hippocampus is necessary for accurate cache retrieval in food-storing birds and is much larger in food-storing birds than in non-storing passerines. It therefore seems probable that seasonal change in caching and seasonal change in the hippocampus are causally related. The peak in recruitment of new neurons into the hippocampus occurs before birds have completed food storing and cache retrieval for the year and may therefore be associated with spacing caches, encoding the spatial locations of caches, or creating a neuronal architecture involved in the recollection of cache sites. The factors controlling hippocampal plasticity in food-storing birds are not well understood. Photoperiodic manipulations that produce change in food-storing behaviour have no effect on either hippocampal size or neuronal recruitment. Available evidence suggests that changes in hippocampal size and neurogenesis may be a consequence of the behavioural and cognitive involvement of the hippocampus in storing and retrieving food.     

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.     

Prey selection and foraging performance of breeding Great Tits Parus major in relation to food availability

We studied the nestling diet and the foraging performance of Great Tits in relation to prey abundance in the field. Numerous experimental studies present data on foraging decisions in captive Great Tits. Little is, however, known about prey selection in the field in relation to the food available and the consequences this has for the food delivery rate to nestlings. Since the foraging performance of the parents is one of the main determinants of fledging weight and juvenile survival, foraging behaviour is an important part of Great Tit reproduction. During the early breeding season up to 75% of the prey biomass delivered to the nestlings were spiders, which is in contrast with other studies. Only when caterpillars reached a size of 10–12 mg (approximately the average size of the spiders caught at that time) did the Great Tits change their preferences and 80–90% of the delivered prey masses were caterpillars, as reported by other authors. This 'switching' between prey occurred within a few days. It was not related to the changes in abundance but to size of caterpillars. The rate at which caterpillars were delivered to the nestlings (in mg/nestling/h) was strongly correlated with the caterpillar biomass available (in mg/m of branches) and nestling growth rate was significantly influenced by the mass of available caterpillars. The results provide evidence why perfect timing of breeding is so important for the Great Tit, and contribute to the understanding of the causal link between food supply, growth and breeding success.     

Evidence of Social Influences on Cache‐Making by Grey Squirrels (Sciurus carolinensis)

Many studies have found that scatter‐hoarding animals change their behaviour when storing food in the presence of conspecifics to minimize the likelihood that their caches will be pilfered; they refrain from caching, move away from conspecifics or choose visually obscured sites. This study reports the first evidence that the presence of conspecifics continues to influence the caching behaviour of a scatter‐hoarding mammal, the grey squirrel, after a suitable cache site has been selected and the hoarder is filling and covering its cache. Wild grey squirrels were filmed when storing preferred and less preferred nuts and when they were alone or with conspecifics present. In line with previous findings, squirrels spent longer travelling from the nut patch and were more vigilant when conspecifics were present. However, squirrels also spent longer disguising their caches and were more likely to stop digging and become vigilant when conspecifics were present than when they were alone. In particular, they were most likely to curtail their digging when storing their preferred nuts in the presence of conspecifics. The results indicate that caching squirrels remain sensitive to the presence of conspecifics until the cache is complete and that they respond flexibly to conspecifics according to the type of food they are storing.     

Breeding time,food supply and fitness components of Blue Tits Parus caeruleus in Mediterranean habitats

We analysed the relationship between the timing of food availability and within-season variation of both reproductive success and nestling body size of Blue Tits Parus caeruleus in Mediterranean habitats. Synchronization between food supply and reproduction was expected to be positively related to fitness components. We measured deviation from maximum food supply using a parameter that we called “time-lag”, which quantifies the degree of synchronization between the date of maximum food requirements by the nestlings and the date of maximum caterpillar supply in the habitat. This parameter was expected to be related to reproductive success as measured by the number and body-condition of fledglings. The predictions were that time-lag should be negatively correlated with the proportion of nestlings raised to fledging and the size of the fledglings. These predictions have been tested in different types of habitat. The results demonstrate that caterpillar supply during a critical nestling period can have a strong influence on fitness components. As predicted, synchronization with caterpillar supply is positively related to the number and body size of fledglings. Since there is large between-habitat variation in the timing of food supply, the key issue seems to be the adjustment to local patterns of food availability.     

Advanced breeding dates in relation to recent climate warming in a Mediterranean montane population of Blue Tits Cyanistes caeruleus

Trends in the onset of breeding, clutch size and numbers of hatchlings and fledglings are examined for a Mediterranean montane population of Blue Tits (Cyanistes caeruleus) subject to recent warming in springtime monitored during 20 years. Blue Tits advanced their breeding dates in relation to mean air temperatures in April and, as a consequence, laid larger clutches. However, increases in the numbers of hatchlings and fledged young over time were not statistically significant after accounting for variables of influence. The entire breeding season seems to have been displaced towards earlier dates by adjusting breeding time to increased temperatures in prebreeding time, to which Blue Tits have been more responsive than Pied Flycatchers (Ficedula hypoleuca) in the same area. The alternative hypothesis, that interference competition with Pied Flycatchers for nestboxes and caterpillars, the main common food base of nestlings, has been the driving force behind the advancement of laying of the Blue Tit population, was not supported. However, the significant advance of breeding dates in Blue Tits has not been sufficient to overcome the precipitous decline in reproductive fitness with the advancement of the season.     

Parental provisioning behaviour in Pied Flycatchers Ficedula hypoleuca is well adjusted to local conditions in a mosaic of deciduous and coniferous habitat

Capsule Pied Flycatchers are better able than Great Tits to adjust their feeding behaviour to varying conditions in the same area.

Aims Great Tits breeding in a mosaic of deciduous and coniferous forests in the northern temperate region exhibit consistently lower breeding success in their preferred deciduous habitat than in coniferous habitat. This was explained by the unexpectedly poor nestling feeding conditions in deciduous forests of this region. We studied whether the same paradox applies to Pied Flycatchers that occupy the same habitats in the same area.

Methods Parental provisioning behaviour was studied using video‐recording and experimental manipulation. Caterpillar abundance and basic breeding parameters were measured in different habitat types.

Results Parental provisioning frequency and the proportion of caterpillars in nestling diet was lower, while food objects were on average larger, in coniferous compared with deciduous habitat. However, the total volume of caterpillars and adult Lepidoptera delivered to nestlings did not differ between habitats. In contrast to Great Tits, offspring body parameters in Pied Flycatchers did not differ between habitat types.

Conclusions These results demonstrate how the relative suitability of particular habitat types varies between species and is dependent upon geographical location.     

Does supplementary calcium reduce the cost of reproduction in the Pied Flycatcher Ficedula hypoleuca?

Studies in acidified as well as in naturally base-poor areas have recently revealed that availability of extra calcium-rich food items is an important component of habitat quality affecting breeding performance in several bird species. However, these mostly short-term studies have provided equivocal results concerning the exact consequences of calcium shortage on different species in different regions. We studied the effect of calcium availability on reproduction of the Pied Flycatcher Ficedula hypoleuca breeding in pine forests in Estonia, NE Europe, over a period of 4 years. Experimental pairs were provided with supplementary calcium-rich material when breeding, while control pairs were left unsupplemented. Experimental females laid larger eggs and their nestlings had longer tarsi than those of controls. Moreover, the mass and condition of females tending larger than average clutches were increased by calcium-supplementation. Our results provide the first experimental evidence that calcium availability may affect the overall cost of reproduction in free-living passerines. We compared these results with similar data for the Great Tit Parus major , collected from the same area during the same study period. Great Tits responded to low calcium availability mainly by restrained reproductive behaviour and reduced breeding success, while Pied Flycatchers invested significantly more in current reproductive effort despite the increased cost of reproduction. Thus, the effects of calcium deficiency on birds seem to be species-specific or population-specific. This partly explains discrepancies between the results of earlier studies.