Stabilizing selection on blue tit fledgling mass in the presence of sparrowhawks

Like British great tits, Belgian blue tits have a lower winter body mass when sparrowhawks are present. Since body mass affects manoeuvrability in small birds, tits may balance the risks of starvation and the risk of hawk predation by varying the amount of extra fat carried during winter. Predation pressure by sparrowhawks on young and inexperienced fledglings is at least as intense as that on the adults during winter. We therefore expected that tit fledgling body mass could also be reduced in the presence of sparrowhawks. In the years after one pair of sparrowhawks settled in a study plot, the mean body mass of blue tit fledglings was lower compared with that in years when there were no sparrowhawks. Furthermore, the shape of the curve relating juvenile survival to fledging mass changed, because the survival of the heaviest fledglings was reduced, which altered the selection differential of juvenile survival as a function of body mass from directional to stabilizing. Of seven published studies on the fledgling body mass–survival relation in tits, all three of the studies conducted in the absence of sparrowhawks showed the highest survival rates for the heaviest young, whereas in all four studies with sparrowhawks present this was no longer the case.     

The effects of hatching date on timing of autumn migration in partial migrants – an individual approach

Timing of spring migration and breeding and their interaction with climate change has been widely studied in recent years, but the possible changes in timing of autumn migration have gained less attention. This work focuses on autumn migration and provides the first multi‐species individual‐based study of how hatching date affects the autumn migration date and migration age by using nestling ring data and re‐trappings of the same individuals during the autumn migration at the Hanko Bird Observatory, Finland. We studied three potentially multibrooded passerines (great tit, blue tit and coal tit) and two single‐brooded birds of prey (goshawk, sparrowhawk), all partially migratory short‐distance migrants. Individuals from late broods migrated at a younger age in all tit species and also in hawks the late hatched individuals tended to migrate at a younger age than the early‐hatched individuals. Late‐hatched individuals migrated later than early‐hatched individuals in blue and coal tits, where the latest hatchers represented second brood individuals. Based on our results, the time from hatching to autumn migration is not constant even among individuals of the same population. Our study indicates that climate warming induced advancement of avian breeding may cause changes in the timing of autumn migration through the frequency of second broods.     

Black-and-white plumage in male pied flycatchers (Ficedula hypoleuca) reduces the risk of predation from sparrowhawks (Accipiter nisus) during the breeding season

Bright colors in birds might signal that they are undesirableas prey (aposematic), an idea that has been difficult to test.When stuffed pied flycatchers Ficedula hypoleuca are exposedto migrating sparrowhawks Accipiter nisus in spring or in autumn,the hawks attack cryptic females more often than bright males.To achieve better statistical control and to assess whethermale plumage also reduces predation risk in the breeding seasonand in the nesting habitat of the pied flycatcher, I placedpairs of male and female flycatcher mounts in similar positionsnear 22 nests of sparrowhawks. The hawks attacked mainly femalemounts, verifying that the preference is real. The sparrowhawkscaught at least 19 live pied flycatchers; 12 young, 5 adultmales, 1 adult female, and 1 female or young. Hawks that caughtan adult male seemed to prefer attacking female mounts. I discussthree interpretations of these results, suggesting that black-and-whitemale flycatchers may benefit from being a novel and aberrantprey, at least early in the breeding season.     

Great tits, Parus major, lay too many eggs: experimental evidence in mid-boreal habitats

This study shows that great tits lay too large clutches in mid‐boreal habitats. First, breeding success, measured with number of fledglings or proportion of eggs that produce fledglings, in northern Finland (65°N) is much poorer than in central and western Europe. Second, brood size manipulations (ca ±30% of the natural mean) revealed that reduced broods produced equal numbers of and larger‐sized fledglings than control and enlarged broods, giving thus the best fitness value for reduced broods. Third, parents of enlarged broods could not adjust (i.e. increase) their feeding effort to the greater number of nestlings. Fourth, extra feeding (about 1/3 of the theoretical maximal needs of the nestlings) during the nestling period resulted in more numerous and larger‐sized fledglings in comparison to control broods. We suggest that the ultimate explanation for the too large clutches is gene flow from the southern population, which prevents local adaptations in the north. Consequently, the main reason for food limitation during the nestling period is that northern great tits apply “southern” decision rules for timing of breeding, clutch size and foraging behaviour. Thus, they tend to breed too early in comparison to the food abundance peak, lay too large clutches in comparison to the level of resources and, perhaps, forage on a too narrow diet (75% caterpillars). Since the late broods that matched the local food abundance peak did not succeed better than the mismatched earlier ones, the most crucial fault of northern great tits seems to be that they overestimate food abundance during peak demands and lay too large clutches. Another explanation for this could be that northern great tits have adopted a brood reduction strategy. However, the long‐term data reveal that years of high breeding success, which would maintain large clutches in the population, are very rare in the north. Therefore, it is unlikely that a brood reduction strategy per se could explain the phenomenon. Instead, it could work together with the gene flow against local adaptation for clutch adjustment.     

Predation by sparrowhawks decreases with increased breeding density in a songbird,the great tit

Predators may regulate prey populations if predation rate increases with prey density. Alternatively, if space-limited (e.g. territorial) predators become satiated when prey exceed a certain density, increased prey abundance may lead to reduced predation rate. These alternatives have been difficult to test experimentally for mobile prey in the wild. We present such a test, manipulating the density of great tits (Parus major) by adding nest boxes in territories of sparrowhawks (Accipiter nisus). Predation rate was measured for young tits after they left the nests. Although the great tit is an important prey, there was no evidence for regulation during the breeding season: the rate of hawk predation declined with increasing density of tits. This result was not confounded by changes in breeding density of alternative prey species (other songbirds). Hawk predation can therefore favour dense breeding in a territorial (solitary) bird, and conspecific attraction and aggregation reported in several territorial species may partly result from predation pressure. This result also has potential implications for conservation work.     

Predation by sparrowhawks Accipiter nisus on male and female pied flycatchers Ficedula hypoleuca in relation to their breeding behaviour and foraging

Bright plumage, song display, and aggressive resource defence in males may cause higher predation on males than on females during the breeding season. However, in birds, higher predation on females is sometimes observed. Parental investment may be high in females (egg-laying, incubation and feeding of offspring), which might lead to a high risk of predation. We studied predation by sparrowhawks Accipiter nisus in relation to behaviour in pied flycatchers Ficedula hypoleuca where breeding males are more conspicuous than females in plumage and behaviour. Male pied flycatchers generally occupied more exposed perches than females. Females were more mobile and foraged more than males, especially prior to and during incubation. During the incubation and nestling stages, when predation on the sexes could be directly compared, sparrowhawks took about the same number of male and female pied flycatchers. During incubation, however, females spent about 77% of the day in the nest and were 4.7 times more vulnerable than males per unit of time available (i.e. outside the nest). A comparison with the chaffinch Fringilla coelebs , where hawks took more females than males, indicates that timing of breeding, foraging behaviour and parental roles of males and females affect predation risk.     

Effect of Japanese lesser sparrowhawks Accipiter gularis on the nest site selection of azure-winged magpies Cyanopica cyana through their nest defending behavior

The frequency of azure-winged magpies Cyanopica cyana that nest in association with breeding Japanese lesser sparrowhawks Accipiter gularis was compared between the 1990s and 2000s. During 1990–1994 azure-winged magpies nested within 50 m of their nests in all of the hawk nest sites studied. During 2005–2006, however, the magpies nested in only two of nine hawk nest sites. During the 1990s Japanese lesser sparrowhawks chased jungle crows away when they intruded within 50 m of their nest. During the 2000s, on the other hand, the hawks attempted to expel few crows at a distance of more than 10 m from their nest. As a result of it, the predation rate of simulated magpie nests located in the hawk nest sites was significantly higher during the 2000s. This suggests that the magpies have the ability to assess the defending behavior of the hawks and judge whether they can rely on their defense. Most azure-winged magpies may stop nesting in association with breeding Japanese lesser sparrowhawks because of the reduced defending range of their nest sites.     

Cuckoo–hawk mimicry? An experimental test

The similarity between many Old World parasitic cuckoos (Cuculinae) and Accipiter hawks, in size, shape and plumage, has been noted since ancient times. In particular, hawk-like underpart barring is more prevalent in parasitic than in non-parasitic cuckoos. Cuckoo-hawk resemblance may reflect convergent evolution of cryptic plumage that reduces detection by hosts and prey, or evolved mimicry of hawks by parasitic cuckoos, either for protection against hawk attacks or to facilitate brood parasitism by influencing host behaviour. Here, we provide the first evidence that some small birds respond to common cuckoos Cuculus canorus as if they were sparrowhawks Accipiter nisus. Great tits and blue tits were equally alarmed and reduced attendance at feeders during and after the presentation of mounted specimens of common cuckoos and sparrowhawks, but not in response to control presentations of collared doves or teal. Plumage manipulations revealed that the strong alarm response to cuckoos depended on their resemblance to hawks; cuckoos with barred underparts were treated like hawks, while those with unbarred underparts were treated like doves. However, barring was not the only feature inducing alarm because tits showed similarly strong alarm to barred and unbarred hawks, and little alarm to barred doves. These responses of tits, unsuitable as hosts and hence with no history of cuckoo parasitism, suggest that naive small birds can mistake cuckoos for hawks. Thus, any cuckoo-hawk discrimination by host species is likely to be an evolved response to brood parasitism.     

BREEDING OF THE GREY WARBLER GERYGONE IGATA AT KAIKOURA, NEW ZEALAND

I studied the breeding of Grey Warblers Gerygone igata (Muscicapidae: Acanthizinae) in forest near Kaikoura, New Zealand, between 1976 and 1979. Only males sang and singing occurred all year. From late July to January pairs defended self-contained territories of 0·25–1·73 ha but they occupied larger home ranges when not breeding. Territorial adults were strictly sedentary all year. The average annual mortality of breeding adults was 18·5% and the predicted life-expectancy 4·9 years, which is remarkable in a bird weighing 6–7 g. The breeding season from first building to last fledging was six months long and it began early. Exceptionally, Grey Warblers may build and lay before the shortest day. As the season progressed warblers nested lower on average, both in absolute terms and relative to the tree nested in and canopy at the site. Warblers built in 7–27 days then delayed up to eight days before laying. Only females built and at no stage of breeding did males feed their mates. Both sexes fed the young. Grey Warblers laid for 15–16 weeks of the year and first clutches were laid asynchronously during 5–6 weeks. Eggs of a clutch appeared at two-day intervals and each egg weighed 1·5 g when fresh (23% of mean adult weight). Clutch size was nearly constant (mean 3·9, mode 4, range 3–5). The incubation period was 17–21 days (mean 19·5 days) and the nestling period 15–19 days (mean 17·2 days). On average the clutch hatched over 1·4 days, even though incubation commenced with the laying of the last egg. Nestlings reached maximum weight on Days 13–14 on average and then receded in weight by 4%, apparently through loss of water. All healthy nestlings exceeded mean adult weight during development by up to 39%. Nestlings from broods of two were at first lighter on average than those from larger broods, but in the second half of the nestling period twins were significantly the heaviest. Grey Warblers were fed for 28–35 days after fledging and they survived well while dependent on parents. Fledglings dispersed up to 3 km or more at independence and only 5% per annum joined the breeding population. Of nests that received eggs, 42% produced at least one fledgling. On average each breeding adult raised 2·0 fledglings per season. Of 265 eggs in 73 nests 70% hatched and 38% produced fledglings. Of 185 nestlings 54% fledged. Probably the main cause of mortality of eggs and nestlings was predation by introduced rodents and mustelids. Grey Warblers raise two small broods slowly during a long breeding season, rather than investing in one large quickly-reared brood. In New Zealand's mild climate the warbler's food supply may not decline severely in winter, and the population of warblers may remain so close to the limit set by food that extra for breeding is hard to obtain. Thus the breeding strategy may be adapted to a restricted food supply.     

Breeding ecology of the Clay-coloured Robin Turdus grayi in lowland Panama

The breeding ecology of Clay-coloured Robins Turdus grayi was studied in 1979 in the former Canal Zone of lowland Panama. Two study areas were chosen close together, Summit Gardens (27-3 ha), an ecological island of habitat suitable for Clay-coloured Robin surrounded by forest, and Morgan's Gardens (3–6 ha), a similar island surrounded by deforested areas and settlements. Eighty-three active nests were found and checked. The breeding season lasted from 2 March to 21 June. In Morgan's Gardens the first broods were raised in the dry season and second broods in the rainy season; in Summit Gardens most broods started at the beginning of the rainy season. In comparison to the closely related Blackbird T. merula in Poland, Clay-coloured Robins build nests on isolated trees or bushes, usually in more conspicuous sites (interpreted as an anti-predator adaptation) and on flexible, horizontal branches off the main trunk. Over 45% of broods were destroyed by predators but other factors causing brood loss were negligible. Nests built on palms were considerably safer. In the dry season the predation rate was low (5%) and increased during the course of the rainy season. Unlike Blackbirds, Clay-coloured Robin nestlings were given a lot of fruit; their diet was diversified. Starvation occurred in 60% of nests although usually only the youngest nestling died. Breeding density in Morgan's Gardens at 50 pairs per 10 ha was much higher than in Summit Gardens at 15>8 pairs per 10 ha. Breeding losses were lower, the nestlings were fed more fruit, the average nestling weight was lower and the production of fledglings per breeding pair was twice as high in Morgan's Gardens. It was concluded that a strategy of settling in a rather overpopulated place with hard foraging conditions but lower predation was better than settling in a area with a better food supply but higher predation.     

Direct and indirect effects of an insect outbreak increase the reproductive output for an avian insectivore and nest‐cavity excavator,the red‐breasted nuthatch Sitta canadensis

Community‐wide food pulses may ameliorate food constraints but may also result in increased competition for other resources and predation rates. In cavity‐nesting vertebrate communities, where the availability of tree cavities can limit reproduction and the reuse of cavities can increase nest predation by squirrels, excavators may maximize their fecundity by creating new cavities in competitor‐ and predator‐rich habitats that undergo food pulses. The reproductive cost associated with excavation (i.e. increased energy allocation early in the breeding season that often delays laying and thereby reduces clutch size), may be reduced if food pulses allow for a longer breeding season and larger clutches. A large‐scale mountain pine beetle Dendroctonus ponderosae outbreak that occurred during our long‐term study (1995–2009) provided a natural food supplementation experiment across 27 sites in British Columbia, Canada. We examined the effects of a reduction in food constraints accompanied with increases in excavation rates, conspecific density and nest predation risk on the fecundity of a facultative excavator, the red‐breasted nuthatch Sitta canadensis. We found a total of 420 nests in tree cavities. Nuthatch clutch sizes ranged from two to nine eggs, and broods from one to nine fledglings per nest. Later clutches were larger at sites and in years with high beetle abundance (mean clutch size of six eggs did not decline later in the season), second broods were produced in outbreak years (usually only one nesting attempt/normal year), and the number of fledglings per successful nest increased with increasing beetle abundance and nuthatch densities, but declined with increased squirrel densities. Since fecundity did not differ between new and reused cavities, the costs and benefits of excavation versus cavity reuse may be neutralized for nuthatches during strong resource pulses. Overall, the beetle outbreak reduced food constraints for nuthatches and provided alternate food for nest predators, allowing increased annual fecundity.     

Alarm call‐based discrimination between common cuckoo and Eurasian sparrowhawk in a Chinese population of great tits

Morphological resemblance of the common cuckoo Cuculus canorus to the Eurasian sparrowhawk Accipiter nisus has been regarded as an example of predator mimicry. Common hosts could distinguish parasites as the result of coevolution, while rare hosts or non‐hosts may mistake cuckoos for hawks because rare hosts or non‐hosts behave similarly when faced with these two species. Birds usually produce alarm calls in addition to showing behavioral responses when in danger. However, previous studies of identification by rare hosts or non‐hosts of sparrowhawks usually lacked experimental evidence of alarm calls. Great tits Parus major, a rare cuckoo host, perform similar behaviors and usually produce alarm calls in response to sparrowhawks and common cuckoos. Here, we tested whether great tits could distinguish common cuckoo from sparrowhawk based on analysis of their alarm calls and the effects of playback of alarm calls on conspecific behavior. Previous studies showed that great tits have a complex communication system that conveys information about predators, and they could perform different kinds of response behavior to different alarm calls. If great tits have not made the ability to distinguish between common cuckoo and sparrowhawk, then their acoustic responses to these two species and their response behaviors in playback experiments should be similar. Specimens of a common cuckoo (parasite), a sparrowhawk (predator) and an Oriental turtle dove Streptopelia orientalis (harmless control) were used to elicit and subsequently record the response behavior and alarm calls of great tits. There was no significant difference in behavioral response among great tits when exposed to the dummy of cuckoo, sparrowhawk and dove. In contrast, they differed significantly in alarm calls. Great tits produced more notes per call that contained increasing D‐type and decreasing I‐type notes when responding to sparrowhawk as compared to cuckoo or dove. In playback experiments, we found that great tits responded more strongly to great tit hawk than to great tit cuckoo or great tit dove alarm calls. Our study suggests that great tits are able to distinguish sparrowhawks from common cuckoos and convey relevant information in alarm calls by adjusting the number and combinations of notes of a single call type.     

Optimal brood size and its limiting factors in the Rufous Turtle DoveStreptopelia orientalis

Optimal brood size and its limiting factors of the Rufous Turtle Dove,Streptopelia orientalis, were studied at the campus of the University of Tsukuba, Japan, during the breeding season in 1990–92. The dove usually lays two eggs in a nest. I made nests of a brood size of one and three by transferring a hatchling from one nest to the other, and compared their fledging success, factors of breeding failure, weight and tarsus length at fledging, growth rate and nestling period with those of a brood of two. The index of fitness (fledgling weight multiplied by average number of fledglings per nest) was almost the same in broods of two and three. However, the highest variation in fledging weight within the brood and the extension of nestling period were observed in broods of three, which caused the extension of inter-brood interval and consequently the smaller number of broods in the total breeding season. Therefore, broods of three would not have an advantage in producing more offspring than broods of two. Crop milk production had an effect on the growth of nestlings in the early phase of the nestling period, but the rapid growth in the granivorous phase compensated for the growth delay of the smallest nestling in broods of three. Small brood size and a large number of broods in a season would also be more effective under high predation pressure.     

ENVIRONMENTAL INFLUENCES ON GROWTH AND SURVIVAL OF NESTLING HOUSE MARTINS DELICHON URBICA

Growth of nestling House Martins was studied in relation to (a) conditions in the external environment and (b) aspects of their breeding biology. The dependence of growth performance on (1) hatchling weights, (2) relative difference in hatchling weights within broods, (3) brood size,(4) season, (5) earliness of breeding in relation to other pairs in the colony, (6) timing of breeding in relation to the median breeding week of the colony and (7) aerial food abundance, was investigated by step-down multiple regression analysis. Up to the stage of the peak brood weight, early laying, small brood sizes and high hatchling weights were associated with higher nestling growth rates. Large relative differences in hatchling weights however tended to depress mean brood weights and increase weight differences (= size hierarchies) within broods. These differences in hatchling weights were considered to contribute significantly to 23% of all nestling deaths, because small, late hatching nestlings suffered very high mortality even when food was abundant. The nestlings which died showed a progressive reduction on growth rates and all succumbed before the 11th nestling day. Because these differences in hatchling weights can be linked to the food supply during laying rather than immediately prior to their death, it is considered that the mortality of these nestlings can ultimately be attributed to the low quality of eggs from which they hatched. There was a tendency for pre-hatching factors to diminish in importance throughout growth, while post-hatching factors increased in importance and, with one exception, were responsible for explaining all the significant variance in the growth characteristics of fledglings. The exception was that differences in wing-lengths in broods could be linked with weight differences at hatching. Food shortages lowered average brood weights prior to fledging. Because pairs breeding during the median breeding week had lighter young, it was inferred that competition for food during this peak of breeding activity had the effect of lowering nestling growth performance, although the overall effect was considered to be small. Early breeding pairs tended to have larger broods, and these large broods showed a lowered growth performance. However, early breeding pairs had relatively smaller weight and wing-length differences, in broods of a given size, than occurred in broods of late breeders. It was therefore concluded that early breeding pairs had some attribute which tended to minimize certain disadvantages of large broods. This effect appeared to be linked to the pair, rather than to season or food supply.     

Parental care in Semipalmated Sandpipers Calidris pusilla: brood desertion by females

Although Semipalmated Sandpipers Calidris pusilla are monogamous, with biparental incubation, most females (86–97%) deserted their broods to the care of their mates, 0–11 days (average 6) after their eggs hatched. Males left the brood an average of 8 days later, shortly before or after the chicks fledged. In several instances, females that deserted in one year remained with the chicks the next year, and vice versa. Females deserted chicks at nests that hatched later in the season at an earlier age than those had hatched earlier ( r ²> o.6). Since females appeared to have an energy deficit in at least some years, and suffered higher mortality rates than males during breeding, it is possible that females deserted broods in order to take advantage of better feeding conditions at migratory stopovers in northeastern North America early in the season. There was little evidence of higher nesting success or earlier hatching date in reuniting pairs, although if both members of a pair returned to the breeding area, 80% reunited. Increased survival of their mate may be most advantageous to males in ensuring that they obtain a female the following year.     

Sex ratio variation among broods of great reed warblers Acrocephalus arundinaceus

The sex of 746 great reed warbler fledglings (from 175 broods) was determined by the use of single primer polymerase chain reaction. The reliability of the technique was confirmed as 104 of the fledglings were subsequently recorded as adults of known sex. The overall sex ratio did not differ from unity. Variation in sex ratios between broods was larger than expected from a binomial distribution. Female identity explained some of the variation of brood sex ratio indicating that certain females consistently produced sex ratios that departed from the average value in the population. The theory of sex allocation predicts that parents should adjust the sex ratio of their brood to the relative value of sons and daughters and this may vary in relation to the quality of the parents or to the time of breeding. In the great reed warbler, the proportion of sons was not related to time of breeding, or to any of five female variables. Of five male variables, males with early arrival date tended to produce more daughters. The sex ratio of fledglings that were a result of extra-pair fertilizations did not differ from that of legitimate fledglings. Hence, there is currently no evidence of that female great reed warblers invest in a higher proportion of sons when mated with attractive males.     

The effects of food availability and distance to protective cover on the winter foraging behaviour of tits (Aves: Parus)

To maximize fitness, many animals must trade off their need to forage efficiently against their need to avoid predators. We studied such a trade-off in four species of tits (Paridae) in a forest near Oxford, UK. During winter, tits form flocks which increase feeding efficiency and reduce predation risk. These flocks feed extensively on beech (Fagus sylvatica) seeds, the abundance of which may be critical for winter survival. Because these seeds drop to the ground, where birds are exposed to sparrowhawk (Accipiter nisus) attack, tits need to trade off their need to find seeds against the proximity to protective cover, provided by dense clusters of hawthorn (Crataegus spp.). The quality of the beech crop differs markedly between trees and years. During a year of abundant beechmast, most tits searched for seeds close to protective cover. This 'safety-first' strategy precluded visits to superabundant food patches if they were too far from protective cover. Among beech trees near to cover, tits tended to prefer those with high seed density. Tits benefited from foraging under trees with high seed density because this correlated significantly with seed mass per square metre and because mean search times decreased with increasing seed density. Finally, we show experimentally that great tits, Parus major, can discriminate between edible (viable) and inedible (empty) seeds.     

Nest predation reduces benefits to early clutch initiation in northern cardinals Cardinalis cardinalis

Life history theory and empirical studies suggest that early breeding confers higher reproductive success, but the extent to which this advantage can be generalized to human‐dominated systems and across species is less well understood. We studied the fitness consequences of clutch initiation for 181 female northern cardinals Cardinalis cardinalis and 1228 nests in forests within urban and rural landscapes of Ohio, USA between 2004–2007. Cardinals that bred earlier made significantly more nesting attempts, but cumulative number of young fledged was similar to that of later‐breeding individuals. The expected number of fledglings produced per successful nest was unrelated to date and remained ~1.8 fledglings across the season, despite the fact that nest survival rates improved dramatically as the season progressed. Because the probability of resighting breeding individuals in subsequent years was unrelated to first clutch initiation date, we have no evidence that clutch initiation affected adult survival. The absence of a clear benefit to early breeding appears to be a consequence of high rates of nest predation early in the breeding season.     

Differential Begging and Locomotory Behaviour by Early and Late Hatched Nestlings Affecting the Distribution of Food in Asynchronously Hatched Broods of Altricial Birds

Distribution of food to early and late hatched nestlings was studied in asynchronously hatched broods of the great tit Parus major, the blackbird Turdus merula, and the fieldfare T. pilaris. Food distribution is related to the locomotory and begging behaviour and positions in the nest of these nestlings. Late hatched (small) nestlings were found to beg more often per feed than bigger nestlings and move more towards favoured positions in the nest to counteract selective feeding of bigger young. The functional significance of these differences in the behaviour of early and late hatched nestlings are discussed. It is argued that they are adaptive by 1) ensuring that each nestling survives when food supplies are ample, and 2) by mediating an optimal brood reduction when food is insufficient to raise the entire brood. The roles of asynchronous hatching, and selective feeding which follows from differential behaviour of early and late hatched young are discussed in relation to food conditions during the breeding season.     

Hematological health state indices predict local survival in a small passerine bird, the great tit (Parus major)

In birds, it has been shown that reproductive effort may impair parental condition, while the relation of different condition indices to subsequent survival is still poorly understood. In this study, we measured body mass and various hematological condition indices in breeding great tits in relation to local survival. Number and quality of nestlings and the occurrence of second broods, potentially reflecting parents' breeding effort, were also considered in analyses. The great tits, both male and female, that returned the following year had had a higher albumin/globulin ratio, lower plasma globulin concentration, and a lower heterophile/lymphocyte ratio during breeding in the preceding year, compared to those who did not return. Surviving males (but not females) also had had a higher level of circulating lymphocytes, compared to nonsurvivors. There was no correlation between breeding effort and survival. We conclude that better immunological state and lower stress in great tits during breeding were positively related to their survival probability.