Calcium effects on life-history traits in a wild population of the great tit (Parus major): analysis of long-term data at several spatial scales

Calcium is an essential micronutrient for birds during egg formation and for skeletal development in nestlings. Habitat level studies suggest that birds breeding in low-calcium areas may be limited in the size or number of eggs they lay and in the quality of their nestlings. However, as birds forage non-randomly and may travel considerable distances to acquire calcium, describing different breeding environments in terms of their calcium availability is problematic. Here we explore the spatial relationships between 300-fold variation in soil calcium and the life-history traits of ca. 6,000 pairs of great tits breeding in a single continuous woodland over 41 years. Controlling for other habitat differences, we found strong positive associations between soil calcium, clutch size and recruitment at spatial scales of over 300 m from each nestbox, suggesting that females may have been travelling inter-territorially to acquire calcium during egg formation. Soil calcium near each nestbox (mean distance = 58 m) was a strong positive predictor of mean fledgling mass, suggesting that local calcium was more important during nestling stages. We found no effect of soil calcium on lay-date or egg mass. This study is the first to provide evidence that small woodland passerines are limited by calcium availability at several different spatial scales. However, experimental work is necessary to test the causality of these spatial patterns.     

Selection for synchronous breeding in the European starling

Colonial birds often demonstrate considerable breeding synchrony. In southern Sweden the semi-colonial European starling initiated the vast majority of clutches within one week. Laying dates were positively skewed so that many birds initiated clutches at similar dates early in the season. Breeding was further synchronised by a particularly strong clutch-size reduction equivalent to one third of an egg per day during the first part of the breeding season. The decline in clutch size with season also held true for separate age-classes of females, for individual females laying at different times at different years and for individual females laying at different times the same year. Trends in breeding success during nestling rearing were unlikely to explain the high degree of breeding synchrony or the seasonal decline in clutch size; nestling survival and growth were weakly related or unrelated to reproductive timing. In contrast recruitment success of fledged offspring declined sharply with season. Even within the synchronous laying period, defined as clutches initiated during the first week each year, local recruitment success declined. It is suggested that the early seasonal decline is caused by selection for synchronous fledging permitting the immediate formation of flocks after fledging, whereas the late seasonal trends may be caused by either population differences in female quality or deteriorating conditions for raising young.     

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.     

Weight change and the body reserves of female blue tits, Parus caeruleus, during the breeding season

A female blue tit lays a large clutch that is commonly heavier than her own body weight. Body reserves cannot provide for the entire clutch and little is known of the importance of these reserves during the breeding season. This study reports on weight changes in adult female blue tits during the breeding season, observed by the use of electronic balances. In addition, data from dissecting 67 adult female blue tits, at different stages of the breeding cycle, were used to investigate the changes in body weight. There was a rapid increase in body weight during a period of 4–5 days before the first egg was laid, which was explained mostly by the development of the oviduct and the production of an egg on the final day. Body weight remained constant during the laying period, before decreasing by approximately one gram after clutch completion, owing to the absence of an egg and the absorption of the oviduct. Body reserves increased during the day after clutch completion, were maintained throughout the incubation period and were mobilized during the first five days of the nestling period. Breeding female blue tits appear to deposit small body reserves to act as a short-term buffer against adverse conditions or to support the brooding phase.     

Caterpillar abundance in the territory affects the breeding performance of great tit Parus major minor

The effects of caterpillar food supply on the breeding performance of a population of the Japanese great tit Parus major minor were investigated. Since more than 90% of the food items in our study site were caterpillars living on trees, we estimated the food availability using 20 frass traps per hectare. The sampling error of this method was about 10% on average, which was accurate enough to detect differences between territories. Food abundance at laying in each territory affected the timing of egg laying. However, food amount after hatching was correlated with clutch size. No relationship was found between fledgling quality and food availability, probably because the effects of local variation in food abundance could be canceled out by parental effort such as extending the foraging area. There was a significant negative correlation between the length of the nestling period and food availability. We suggest that parent tits decide the timing of fledging at the point where two factors, predation risk before fledging and additional improvement of nestling quality, are balanced. Food availability just after fledging affected the length of post-fledging parental care; it seems that fledglings in “poor” territories would have had difficulty in finding food and hence needed to depend on their parents longer than those in “rich” territories. Received: 10 June 1997 / Accepted: 29 December 1997     

Latitudinal variation in blue tit and great tit nest characteristics indicates environmental adjustment

Aim The laying of eggs and the building of a nest structure to accommodate them are two of the defining characteristics of members of the class Aves. Nest structures vary considerably across avian taxa and for many species the structure of the completed nest can have important consequences both for parents and their offspring. While nest characteristics are expected to vary adaptively in response to environmental conditions, large‐scale spatial variation in nest characteristics has been largely overlooked. Here, we examine the effects of latitudinal variation in spring temperatures on nest characteristics, including insulatory properties, and reproductive success of blue tits, Cyanistes caeruleus, and great tits, Parus major. Location Great Britain. Methods Nests and reproductive data were collected from seven study sites, spread over 5° of latitude. The nest insulatory properties were then determined before the nests were separated into nest base material and cup lining material. Results As spring temperatures increased with decreasing latitude, the mass of the nest base material did not vary in either species, while the mass of the cup lining material and nest insulatory properties decreased in both species. This suggests that in response to increasing temperatures the breeding female reduces the mass of the cup lining material, thereby maintaining an appropriate microclimate for incubating and brooding. The mean first egg date of both species advanced with decreasing latitude and increasing spring temperatures, although clutch size and brood size at hatching and fledging did not vary. Main conclusions This is the first study to demonstrate that the nest‐construction behaviour of birds varies in response to large‐scale spatial variation in ambient temperatures. Therefore, nest composition reliably indicates environmental conditions and we suggest that studies of nest structure may be sentinels for the early signs of rapid climate change.     

Testing the relationship between clutch size and brood size in the Coot (Fulica atra)

The time between egg laying and chick fledging is of crucial importance for the survival of young birds. I analyzed breeding output at consecutive phases of growth of young Coots (Fulica atra) relative to the clutch size and laying date. Considering the specific breeding biology of the Coot, I tested whether chick survival reveals clutch size-dependent variability. Clutch size did not affect hatching success; it only affected brood size, and that merely temporarily. During the first 20 days after hatching, i.e. during the time of the highest chick mortality, birds with larger clutches lost chicks at a higher rate. As a result, the number of fledged chicks was independent of the initial number of chicks, and pairs with different clutch sizes had a similar number of fledglings. The laying date had no effect. This pattern of age-related chick survival points to the greater role of the type of chick growth (semi-precocial) and behavior in their survival.     

Caterpillar biomass depends on temperature and precipitation,but does not affect bird reproduction

Complex changes in phenological events appear as temperatures are increasing: In deciduous forests bud burst, hatching of herbivorous caterpillars, egg laying and nestling time of birds when feeding chicks on caterpillars, may differentially shift into early season and alter synchronization. If timing of bird reproduction has to match with short periods of food availability, phenological mismatch could negatively affect reproductive success. Using a unique empirical approach along an altitudinal temperature gradient, we firstly asked whether besides temperature, also precipitation and leaf phenology interplay and affect caterpillar biomass, since impacts of rainfall on caterpillars have been largely neglected so far. Secondly, we asked whether abundance of caterpillars and thereby body mass of great tit nestlings, which are mainly fed with caterpillars, vary along the altitudinal temperature gradient. We demonstrated that next to temperature also precipitation and leaf phenology affected caterpillar biomass. In our beech forest, even along altitudes, caterpillars were available throughout the great tit breeding season but in highly variable amounts. Our findings revealed that although timing of leaf phenology and great tit breeding season were delayed with decreasing temperature, caterpillars occurred synchronously and were not delayed according to altitude. However, altitude negatively affected caterpillar biomass, but body mass of fledglings at high altitude sites was not affected by lower amounts of caterpillar biomass. This might be partially outweighed by larger territory sizes in great tits.     

The effects of environmental heterogeneity on multivariate selection on reproductive traits in female great tits

Describing natural selection on phenotypic traits under varying environmental conditions is essential for a quantitative assessment of the scale at which adaptation might occur and of the impact of environmental variability on evolution. Here we analyzed patterns of multivariate selection via fecundity and viability on three reproductive traits (laying date, clutch size, and egg weight) in a population of great tits (Parus major). We quantified selection under different environmental conditions using (1) local variation in breeding density and (2) distinct areas of the population's habitat. We found that selection gradients were generally stronger for fecundity than for viability selection. We also found correlational selection acting on the combination of laying date and clutch size; this is the first documented evidence of such selection acting on these two traits in a passerine bird. Our analyses showed that both local breeding density and habitat significantly influenced selection patterns, hence favoring different patterns of reproductive investment at a small-scale relative to typical dispersal distances in this species. Canonical rotation of the nonlinear selection matrices yielded similar conclusions as traditional nonlinear selection analyses, and also showed that the main axes of selection and fitness surfaces varied over space within the population. Our results emphasize the importance of quantifying different forms of selection, and of including variation in environmental conditions at small scales to gain a better understanding of potential evolutionary dynamics in wild populations. This study suggests that the fitness landscape for this species is relatively rugged at scales relevant to the life histories of individual birds and their close relatives.     

Annual and seasonal reproductive trends in the Lesser Spotted Woodpecker Dendrocopos minor

Annual and seasonal variation in reproductive timing and performance were studied in a population of the Lesser Spotted Woodpecker Dendrocopos minor over 10 years in southern Sweden. The median laying date of the first egg varied by up to 17 days between years, being generally larger than the variation of laying dates within years. Neither clutch size, brood size in successful nests, fledging success in successful nests nor mean nestling weight differed significantly between years. There was no trend for mean clutch size to vary between early and late years. In spite of a more than threefold variation in population size, no reproductive variable demonstrated an apparent density-dependence. Within the season, clutch size declined steeply with increasing clutch initiation date, whereas fledging success and nesting success did not, leading to a trend in brood size almost identical to the trend in clutch size. The survival prospects of fledged young declined with increasing clutch initiation date, and it is argued that the clutch size laid is a strategic adjustment to laying date. Out of 124 breeding attempts, 34% did not produce fledged young. In 9% of the breeding attempts, pairs laid no eggs. At least 20% of the breeding attempts failed after egg-laying. The most common cause of breeding failure was loss of the breeding partner followed by nest abandonment (40% of the failures). Only 16–28% of the failures were due to predation on the nest. Most complete failures, and also partial losses from nests, occurred at the early breeding stages. It is argued that the early nestling phase may be a critical stage, which the woodpeckers adjust to coincide with the seasonal food peak, explaining the strikingly late breeding season compared with other non-migrant species.     

Arrival date, age and breeding success in white stork Ciconia ciconia

Early arrival to breeding grounds is a life history trait in birds that can result in fitness benefits. We studied the relationship between arrival date and breeding success of individuals in a central Iberian population of white stork Ciconia ciconia , between 1999 and 2005, and the ways in which other potential factors, such as age or sex, affect this relationship. Our results showed that age was the factor most closely related to arrival date and breeding success. Older individuals returned earlier to the breeding grounds, achieved larger clutch sizes and produced more chicks than younger birds. After controlling statistically for age effect, breeding probability (laid eggs or not) and laying date were still significantly explained by arrival date. A higher probability of failure to reproduce (no eggs laid) was found in birds arriving later than in those arriving early. However, clutch size and nestling success (number of nestlings in the nest 40 days after hatching) were not correlated with arrival date. Food availability in the study area throughout the breeding cycle, due to a nearby rubbish dump, could be the factor mitigating differences in clutch size and nestling success related to individual arrival date.     

Leafing phenology and timing of egg laying in great tits Parus major and blue tits P. caeruleus

To evaluate the importance of tree leafing for the start of laying and clutch size of birds, we compared the breeding phenology of great tits Parus major and blue tits P. caeruleus between one coastal and two inland sites in the same geographical region. Because of the cooling influence of the sea, trees at the coastal site were known to initiate budburst about a week later than at the inland sites. During 5 years, breeding by the tits and the leaf phenology of birch Betula pendula , and oak Quercus robur were monitored. The leaf phenology of birch and oak explained a significant part of the between-year variation in the start of egg laying in blue and great tits, respectively. The tits started laying earlier at the sites with an early budburst, i.e. normally inland. However, leaf phenology was not an absolute cue to the start of laying, since blue tits laid earlier relative to leafing at the inland site than at the coastal site, and both tit species laid eggs earlier relative to leafing during late springs. In neither species was clutch size affected by leafing phenology. However, great tit females at the coastal site consistently produced fewer eggs than did those at the inland site. No such difference was found in the blue tits. Although leafing phenology may predict the start of laying in tits, other factors also influence its timing. These factors might include other cues, or differing life-history trade-offs depending on site or general climatic factors during the spring.     

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.     

Increased survival and breeding performance of double breeders in little penguins Eudyptula minor,New Zealand: evidence for individual bird quality?

The little penguin Eudyptula minor is unique among penguin species in being able to fledge chicks from two clutches in one breeding season. Pairs laying two clutches in a given season make a higher reproductive investment, and may be rewarded by a higher reproductive success as they may raise twice as many chicks as pairs laying one clutch. The higher effort made by pairs laying two clutches could correlate negatively with survival, future reproductive performance or offspring survival, indicating a cost of reproduction. Conversely, a positive relationship between the number of clutches produced in a given breeding season and survival, future reproductive performance or offspring survival would indicate that birds laying two clutches belonged to a category of birds with higher fitness, compared to birds laying only one clutch in the season. In this study we used a long‐term data set taken from an increasing population of little penguins in Otago, SE New Zealand. We modelled the relationship between the number of clutches laid in a breeding season and survival probability, reproductive performance in the next breeding season and first year survival of offspring using capture‐recapture modelling.
Birds laying two clutches produced 1.7 times more fledglings during a breeding season than pairs laying one clutch. We found that birds laying two clutches had a higher probability of breeding in the following breeding season, a higher probability of laying two clutches in the following breeding season and a higher survival probability. There was no overall difference in post‐fledging survival between the young of birds producing one clutch and the young of birds producing two clutches. However, the survival of young of single clutch breeders declined with laying date, whereas the young of double clutch breeders had the same survival rate irrespective of laying date. For a subset of data with birds of known age, we found evidence that the probability of laying two clutches increased with age. However, there were also indications for differences among birds in the tendency to lay two clutches that could not be attributed to age. We tentatively interpret our results as evidence of quality difference among little penguin breeders.     

Geographic variation in reproduction and survival of kelp gulls Larus dominicanus vetula in southern Africa

Different populations of a species tend to vary in survival and reproduction, but the extent and scale of such spatial variation are poorly known. We estimated survival and clutch size of kelp gulls Larus dominicanus vetula across their entire African range. At this large geographic scale, we found no evidence for spatial variation in survival, and there was no variation in clutch size. However, there was considerable variation in clutch size among colonies within regions. Over the whole study, mean annual survival of juvenile and adult birds was 0.44 and 0.84, and mean clutch size was 2.2 eggs. A matrix population model showed that population growth was least sensitive to variation in clutch size, and the observed variation in clutch size could not fully account for the observed variation in population growth among colonies and regions. Our results thus suggest that dispersal and/or variation in survival (including egg/nestling survival) at a small spatial scale are also important for the spatial pattern of kelp gull population dynamics. These results are consistent with a metapopulation approach to spatial population dynamics.     

Density dependent effects between three competitive bird species

Summary Density and breeding success of the great tit Parus major, blue tit Parus caeruleus and collared flycatcher Ficedula albicollis were studied in nest box colony in oak forest over a period of 19 years.Intraspecific density dependent clutch size reduction was found with blue tit and great tit. In interspecific relation the high density of blue tits reduced the clutch size of great tits.In the hatching period neither intraspecific nor interspecific density dependence were showed between the tits when the third competitive species, collared flycatcher was present. The collared flycatcher significantly reduced the hatching success of both tit species and the number of fledglings of great tit.The effects of the great tits and combined density of the great and blue tits on the hatching failure and number of fledglings of collared flycatcher were found when the density of the tits was high. There were not significant relationships to the single density of blue tits.The temporal variability of the competition of the three bird species is discussed.     

Reproductive success and nestling diet in the Lesser Spotted Woodpecker ( Picoides minor ): the early bird gets the caterpillar

Populations of Lesser Spotted Woodpecker (Picoides minor) are decreasing in size, necessitating management strategies. However, data on the reproductive biology of this species are scarce. The aim of this study was to investigate the influence of timing of breeding and nestling diet on the reproductive success of the Lesser Spotted Woodpecker and thereby contribute to an understanding of the determinants influencing its reproductive success. During 6 study years between 1996–2003, we investigated various variables of reproductive success in a Lesser Spotted Woodpecker population in the Taunus low mountain range, Germany. We observed nestling feeding at breeding holes to assess the composition of nestling diet and how it changes during the season. Clutch size, number of fledglings as well as body mass of the nestlings declined with the start of egg laying. Pairs composed of individuals that had breed together in previous years started egg laying earlier than newly established pairs. Nestling diet consisted mainly of aphids, caterpillars, craneflies and wood-living larvae. The composition changed considerably within the breeding season in both early and late broods. However, early broods were provided with caterpillars more frequently, whereas late broods received wood-dwelling larvae more often. Our results suggest that Lesser Spotted Woodpeckers time their breeding so as to coincide with the early nestling stage with the highest availability of caterpillars, since this is the time of highest energy demand. Moreover, food availability declines during the breeding season, and the decline in reproductive performance seems to be an effect of this development.     

The importance of food quantity and quality for reproductive performance in alpine water pipits (Anthus spinoletta)

Studies relating reproduction to food availability are usually restricted to food quantity, but ignore food quality and the effects of habitat structure on obtaining the food. This is particularly true for insectivorous birds. In this study we relate measures of reproductive success, time of reproduction and nestling size of water pipits (Anthus spinoletta) to biomass, taxonomic composition and nutritional content of available food, and to vegetation structure and distance to feeding sites. Clutch size was positively correlated with the proportion of grass at the feeding sites, which facilitates foraging. This suggests that water pipits adapt their clutch size to environmental conditions. Also, pipits started breeding earlier and produced more fledglings when abundant food and a large proportion of grass were available, probably because these conditions allow the birds to gain more energy in less time. The number of fledglings was positively correlated with the energy content of available food. No significant relationships were found between feeding conditions and nestling size or the time that nestlings took to fledge. This suggests that water pipits do not invest more in individual nestlings when food conditions are favourable but rather start breeding earlier and produce more young. Taxonomic composition and nutritional content of prey were not correlated with any of the reproductive parameters, indicating that profitability rather than quality of food affects reproductive success. Received: 31 May 1996/Accepted: 12 August 1996     

Phenological sensitivity to climate change is higher in resident than in migrant bird populations among European cavity breeders

Many organisms adjust their reproductive phenology in response to climate change, but phenological sensitivity to temperature may vary between species. For example, resident and migratory birds have vastly different annual cycles, which can cause differential temperature sensitivity at the breeding grounds, and may affect competitive dynamics. Currently, however, adjustment to climate change in resident and migratory birds have been studied separately or at relatively small geographical scales with varying time series durations and methodologies. Here, we studied differential effects of temperature on resident and migratory birds using the mean egg laying initiation dates from 10 European nest box schemes between 1991 and 2015 that had data on at least one resident tit species and at least one migratory flycatcher species. We found that both tits and flycatchers advanced laying in response to spring warming, but resident tit populations advanced more strongly in relation to temperature increases than migratory flycatchers. These different temperature responses have already led to a divergence in laying dates between tits and flycatchers of on average 0.94 days per decade over the current study period. Interestingly, this divergence was stronger at lower latitudes where the interval between tit and flycatcher phenology is smaller and winter conditions can be considered more favorable for resident birds. This could indicate that phenological adjustment to climate change by flycatchers is increasingly hampered by competition with resident species. Indeed, we found that tit laying date had an additional effect on flycatcher laying date after controlling for temperature, and this effect was strongest in areas with the shortest interval between both species groups. Combined, our results suggest that the differential effect of climate change on species groups with overlapping breeding ecology affects the phenological interval between them, potentially affecting interspecific interactions.     

Habitat fragmentation influences nestling growth in Mediterranean blue and great tits

In patchy forest areas, the size of the forest patch where birds breed has a strong influence on their breeding success. However, the proximate effects contributing to lowering the breeding success in small forest patches remain unclear; and a shortage of crucial resources in those forest patches has been suggested to account in some degree for this failure. With the aim to further investigate this issue, we have monitored the breeding cycle of blue and great tits in three ‘large’ forest patches (ranging between 26.5 and 29.6 ha) and twelve ‘small’ forest patches (ranging between 1.1 and 2.1 ha) in a Mediterranean area in central Spain, during three years (2011–2013). We also recorded the nestling diet inside the nest-boxes with the aid of handy-cams. Only males significantly differed between forest patch size categories; being on average younger and with better body condition in small patches for great and blue tits respectively. Reproductive traits did not vary between forest patch size categories, but the body condition of blue tit nestlings and the size of great tit nestlings did, being significantly better and larger respectively in large forest patches. The recruitment rate of blue tit nestlings was also higher in large patches. Regarding nestling diet, blue tits did not differ but great tits did, delivering a larger amount of caterpillars in large forest patches. Most variation in the reproductive traits occurred between years, probably due to annual differences in environmental conditions. This study suggests that food supply could be limiting the breeding success of birds above all in small patches, but also in large patches under particular environmental conditions.