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.     

Fitness-related consequences of egg mass in nestling house wrens

The fitness-related consequences of egg mass, independent of confounding influences associated with parental quality, remain poorly understood for wild birds in general and for passerines in particular. We performed cross-fostering experiments to test the hypothesis that egg mass, independent of parental quality, is the primary determinant of fitness-related traits in nestling house wrens (Troglodytes aedon), an insectivorous passerine. Nestling mass was significantly correlated with the mass of the eggs from which nestlings hatched early but not late in the nestling period in early-season broods. In contrast, in late-season broods, nestling mass was correlated with egg mass until nestlings achieved asymptotic mass. Neither nestling growth nor survival to nest leaving was related to egg mass in either early- or late-season broods; however, nestlings in late-season broods grew more slowly than did nestlings in early-season broods. We propose that nestling mass and egg mass remained correlated throughout the nestling period in late-season broods because decreased arthropod food resources late in the breeding season constrain parents'' ability to provision nestlings. We conclude that female house wrens in this population trade-off clutch size for greater egg mass to maximise reproductive success in late-season broods.     

Two cooperative breeding cases in Lesser Spotted Woodpecker Dendrocopos minor

During a study of a colour-marked population of Lesser Spotted Woodpecker (LSW) in NE Spain we found two cases of cooperative breeding, in which two females and a male were rearing the brood. One of the females was involved in both cooperative breeding cases, in two consecutive years. We analysed the feeding contribution of each individual to nestling care as well as feeding rates to nestlings. In both cases the male showed a (compensatory) reduction in nestling feeding due to the involvement of the second female. Total feeding rate increased slightly compared to regular (noncooperative) monogamous pairs. The high breeding success found in the second case (six eggs/six nestlings) is thought to be due to the cooperation of the three adults in nestling care. The two cooperative breeding cases were found in territories that suffered from significant selective logging of dead trees. This logging is thought to be a possible anthropogenic example of the ecological constraints hypothesis and a significant factor that could force atypical cooperative breeding attempts. A locally biased sex-ratio might result in the cooperation of two females and a male in both cases. No evidence of kinship was found among members of the cooperative breeding teams. We could not determine whether the cooperative breeding cases were monogamous or polygamous. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Importance of feeding ecology to the reproductive success of Blackbirds Turdus merula nesting in rural habitats

The feeding ecology of Blackbirds Turdus merula breeding in contiguous woodland and farmland habitats was studied over three years. The aim of the study was to investigate how reproductive success was influenced by nestling diet and the provisioning rates of parents feeding nestlings. Parental provisioning rates increased with brood size, and consequently individual nestlings were no lighter in larger broods. None of the environmental factors measured had strong effects on parental provisioning rate. The nestling diet was dominated by caterpillars and earthworms, the former occurring in a short period in the middle of the breeding season. The availability of earthworms was higher in woodland and was dependent on rainfall in farmland. Nestling mass and provisioning rates were marginally higher under predominantly earthworm diets. Nestling mass increased with rainfall in farmland only, and was higher in farmland than in woodland or woodland-edge, although it is doubtful whether this result is of any significance for fledgling survival. Overall, Blackbirds were able to provision their nestlings adequately throughout the breeding season across a range of conditions. There was no evidence to suggest that reproductive success was constrained by aspects of feeding ecology within the natural range of brood size.     

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.     

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 pattern of Dunlin Calidris alpina distribution and abundance in relation to habitat variation in the Flow Country of northern Scotland

Capsule Folivorous caterpillars constituted the majority of nestlings’ food in a primeval forest. Blue Tit broods only partially matched the caterpillar peak, and the mismatch did not affect food composition or nesting success.

Aims To describe factors influencing the timing of reproduction in Blue Tits under primeval conditions (Bia?owie?a National Park, Poland) and to check whether they schedule breeding so as to synchronize broods with a seasonal caterpillar peak.

Methods We gathered information on phenology of leaf development, seasonal availability of folivorous caterpillars (frass collection), timing of Blue Tit breeding, composition of its nestling food, and nest fate over a three-year period.

Results Caterpillars constituted c. 74% of nestling diet, but only 17–65% of broods matched the caterpillar peak in any season. Neither total nest loss, nor frequency of brood reduction depended on the level of mismatch. Caterpillar availability was probably adequate every year, regardless of the amount of mismatch, and no selective advantage of precise matching was detectable. Phenological events at all trophic levels occurred earlier in warmer springs. Egg-laying coincided with tree bud burst and appearance of caterpillars, but was not critically dependent on their timing.

Conclusion The observations are consistent with the view that Blue Tits under primeval conditions in Bia?owie?a National Park, Poland, breed as early as possible, rather than synchronizing their breeding with the caterpillar peak later in the season.     

Sources and timing of calcium intake during reproduction in flycatchers

Calcium availability may limit the reproductive output of birds and snail shells are considered to be the main source of calcium in many passerine species. This study of collared (Ficedula albicollis) and pied (F. hypoleuca) flycatchers evaluates calcium intake of a natural diet in Central Europe, and sex differences in the utilization of experimentally supplemented sources of calcium during the entire breeding period in aviary birds. The study provides the first evidence that successful reproduction of these species depends on the availability of woodlice (Isopoda) and millipedes (Diplopoda). Each of these two components provided about 3 times more calcium than the snail shells contained in a natural nestling diet. The breeding performance of aviary birds was poor when only snail shells and the fragments of eggshells were provided in food, i.e., irregular laying, smaller clutches, eggshell defects (25 of 53 eggs), and eggs dried-up during incubation. In contrast, no defective eggshell or dried-up eggs were found and the overall breeding performance increased 2–3 times when woodlice were added to the food. Females increased their intake of woodlice during both the pre-laying and laying periods, and both sexes did so during the nestling period. Both sexes took more woodlice in the evening than in the morning, independent of the nesting stage. Intake by females was low until 4 days before laying the first egg, then increased to the highest level, dropping immediately after laying the last egg. Intake of woodlice by both sexes increased steadily from hatching until the nestling age of about 10–12 days decreasing thereafter, which corresponds with the period of rapid skeletal growth. In contrast, the intake of mealworms increased until the nestling age of 13–14 days leveling off thereafter which corresponds with the growth curve of nestling body mass.Due to an error in the citation line, this revised PDF (published in December 2003) deviates from the printed version, and is the correct and authoritative version of the paper.     

Consequences of a changing environment on the breeding phenology and reproductive success components in a long-distance migratory bird

Migratory birds have a narrow time window to breed, especially in the Arctic, where early nesting typically yields the highest reproductive success. We assessed temporal changes (1991–2015) in reproductive success components in relation to timing of breeding in greater snow geese (Chen caerulescens atlantica). This species breeds in the Canadian Arctic, a region that has experienced a strong warming trend. We tested the effect of laying or hatching date, year and their interaction on six reproductive components: Total clutch laid, nesting success, egg survival, hatching success, prefledging, and postfledging survival. Over 25 years, mean laying date changed little, even though it advanced 1.8 days in early breeders and was delayed 3 days in late breeders. Likewise, the number of eggs in nests initiated early in the season decreased by 0.6 egg, whereas in late nests it increased by 0.3 egg. Success of nests initiated early and late in the season was lower than nests initiated near the population mean, and consistently increased over time. The proportion of eggs surviving to partial predation and postfledging survival decreased with laying date but the pattern did not change over time. In contrast, prefledging survival was not affected by laying date initially but declined in nests initiated late in the season toward the end of the study. Overall, nests initiated close to the population mean showed little temporal change for most components of reproductive success and seem to be less affected by environmental change than nests initiated early and late in the season.     

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.     

Habitat quality, nestling diet, and provisioning behaviour in great tits Parus major

Variation in early nutrition is often a strong predictor of offspring condition and fitness. In the case of woodland passerine birds, nestling diet is determined by the spatiotemporal distribution of prey items such as caterpillars during the nestling period, and is usually quantified as differences in provisioning behaviour between habitats. However, the habitat level does not account for variation between individual territories, the level at which competition and selection are assumed to operate. Here we use nestbox cameras and Radio Frequency Identification technology (RFID) to simultaneously assess variation in both nestling diet (components) and provisioning rates (quantity) among a sample (n=22) of different quality great tit Parus major territories selected from a larger breeding population (n=310 fledged broods) in a single year. Caterpillars were by far the most numerous item provisioned to nestlings (mean=75% of prey items), as expected given the known importance of this food source for this species. Broods raised close to an oak tree, or far from the woodland edge, were provisioned the highest proportion of caterpillars. Provisioning rates declined seasonally and there was a weak association between low provisioning rates and caterpillar rich diets. During the first week of the nestling stage, nestling condition was unrelated to the proportion of caterpillars in the diet, provisioning rates and oak proximity. Condition at fledging was slightly improved in broods fed a higher proportion of caterpillars in the diet and in broods raised close to an oak tree. However, in our data early breeding was the only predictor of recruitment success, although power was low for this test. Analyses of long‐term data (41 years) from the same population confirmed a relationship between oak proximity and fledgling mass, but not recruitment success. Our results suggest that territory level environmental variation can affect offspring condition, probably through observed changes in nestling diet, but that such variation does not necessarily produce discernable effects on offspring fitness.     

Seasonal decline in provisioning effort and nestling mass of Blue Tits Cyanistes caeruleus: experimental support for the parent quality hypothesis

A common feature of many birds breeding in seasonal environments is that fitness‐related parameters such as nestling mass or survival decline as the breeding season progresses. Consequently, there is a tendency for early breeders to have better reproductive performance than individuals breeding later in the season. This variation could be caused by factors associated with the date of laying, such as changing environmental conditions (date hypothesis), or by differences in parental quality between early and late breeders (parent quality hypothesis). To evaluate the relative importance of both hypotheses, we manipulated hatch dates of Blue Tits Cyanistes caeruleus by exchanging clutches with different incubation stages and assessed the impact on nestling mass, nestling diet and provisioning rates. Mean nestling mass declined significantly over the season. This was the combined result of differences in parental quality, which dominated in the early part of the season, and the influence of hatching date (date effect per se), which prevailed later in the season. Nestling diet composition was apparently uninfluenced by the manipulation, suggesting that deteriorating food supplies are the primary reason for the seasonal variation in the nestling diet. Counter to the date hypothesis, delayed parents did not feed their young less than control pairs did, but in fact exhibited higher provisioning rates. Our results suggest that in this population, parental quality seems to constrain post‐hatching reproductive performance and such intrinsic limitations may help to explain why certain individuals breed later.     

Parental care of kestrels living in stable and varying environmental conditions

We studied parental behaviour of two populations of Eurasian kestrels (Falco tinnunculus) living in environments with different reproductive constraints: a northern Italy population, resident, with relatively long reproductive season and almost stable prey availability, and a western Finland population, long-distance migrant, with short reproductive season and fluctuating availability of main prey (voles). The Finnish population was studied during a low vole availability year. We aimed to test the supposition that in variable environments the few pairs capable of breeding during low prey availability years show high-quality parental care. To estimate the quality of parental care in both Finnish and Italian pairs we measured the difference in the number of prey delivered by the male and by the female within each pair. The parental care, measured by that parameter, was greater among Italian pairs during both the incubation and early nestling periods. However, this was not the case in the late nestling period, when food requirements of offspring are greater. Moreover, the Finnish and Italian pairs produced about the same number of fledglings. Therefore, our data supported the initial supposition and we conclude that the few Finnish pairs capable of breeding in a low vole availability year show parental care similar to Italian pairs.     

Cascading costs of reproduction in female house wrens induced to lay larger clutches

In many species, females produce fewer offspring than they are capable of rearing, possibly because increases in current reproductive effort come at the expense of a female's own survival and future reproduction. To test this, we induced female house wrens (Troglodytes aedon) to lay more eggs than they normally would and assessed the potential costs of increasing cumulative investment in the three main components of the avian breeding cycle – egg laying, incubation and nestling provisioning. Females with increased clutch sizes reared more offspring in the first brood than controls, but fledged a lower proportion of nestlings. Moreover, nestlings of experimental females were lighter than those of control females as brood size and prefledging mass were negatively correlated. In second broods of the season, when females were not manipulated, experimental females laid the same number of eggs as controls, but experienced an intraseasonal cost through reduced hatchling survival and a lower number of young fledged. Offspring of control and experimental females were equally likely to recruit to the breeding population, although control females produced more recruits per egg laid. The reproductive success of recruits from broods of experimental and control females did not differ. The manipulation also induced interseasonal costs to future reproduction, as experimental females had lower fecundity than controls when breeding at least 2 years after having their reproductive effort experimentally increased. Finally, females producing the modal clutch size of seven eggs in their first broods had the highest lifetime number of fledglings.     

Constraints on, and determinants of, the annual number of breeding attempts in the multi-brooded Black Redstart Phoenicurus ochruros

With reference to models predicting patterns of reproduction in multi-brooded species, I analysed some of the factors potentially affecting the number of breeding attempts per season made by female Black Redstarts Phoenicurus ochruros in the central Swiss Alps. The maximum number of successful broods per female per season was three, the median two. The proportion of females initiating only a single clutch per season varied from 16 to 58% over 10 years. Variation in the frequency of single brooding between years was positively related to the date of the onset of breeding. Egg-laying started after a temperature-related threshold value was crossed in April. Females frequently re-nested before fledglings of the previous brood were independent. Short interbrood intervals were associated with triple brooding. The interbrood interval shortened during the breeding season. Intra-individual variation in the number of breeding attempts per season was partitioned according to female age, laying date of the season's first egg (standardized in relation to the median date of first laying within a year) and the occurrence of breeding failures before the ultimate nesting attempt. However, mortality during the season often terminated breeding early, in particular among novice breeder females. Seasonal reproductive success increased linearly with each additional breeding attempt. The productivity of a breeding attempt was independent of the time in the season. Increased reproductive effort affected neither current or future survival nor reproduction of females. The reproductive patterns and trade-offs in multi-brooded Black Redstarts contrast in many aspects with those found in single-brooded species. They are in accordance with models predicting that multi-brooded species are selected to start breeding as early as possible, continue breeding for as long as conditions are suitable, and save time by overlapping broods.     

Climate change and fitness components of a migratory bird breeding in the Mediterranean region

The increase in spring temperatures in temperate regions over the last two decades has led to an advancing spring phenology, and most resident birds have responded to it by advancing their onset of breeding. The pied flycatcher (Ficedula hypoleuca) is a long‐distance migrant bird with a relatively late onset of breeding with respect to both resident birds and spring phenology in Europe. In the present correlational study, we show that some fitness components of pied flycatchers are suffering from climate change in two of the southernmost European breeding populations. In both montane study areas, temperature during May increased between 1980 and 2000 and an advancement of oak leafing was detected by using the normalized difference vegetation index (NDVI) to assess tree phenology. This might result in an advancement of the peak in availability of caterpillars, the main prey during the nestling stage. Over the past 18 yr, the time of egg laying and clutch size of pied flycatchers were not affected by the increase in spring temperatures in these Mediterranean populations. However, this increase seems to have an adverse effect on the reproductive output of pied flycatchers over the same period. Our data suggest that the mismatch between the timing of peak food supply and nestling demand caused by recent climate change might result in a reduction of parental energy expenditure that is reflected in a reduction of nestling growth and survival of fledged young in our study populations. The data seem to indicate that the breeding season has not shifted and it is the environment that has shifted away from the timing of the pied flycatcher breeding season. Mediterranean pied flycatchers were not able to advance their onset of breeding, probably, because they are constrained by their late arrival date and their restricted high altitude breeding habitat selection near the southern border of their range.     

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.     

A male destroying an egg in a cooperative breeding attempt in Lesser Spotted Woodpecker Dendrocopos minor

A cooperative breeding case was found in Lesser Spotted Woodpecker (LSW) in which a male and three different females were involved. The contributions of each member of the cooperative breeding team were quantified during nest excavation and incubation. This is the first such report on LSW. During daytime, the nest was mainly occupied by the male during the final excavation phase, nest guarding, and egg laying. Aggressive interactions were recorded between two of the females during nest excavation, egg laying and incubation. The male was recorded destroying an egg presumably laid by the dominant female. The highest contribution during daytime incubation was made by the dominant female (39%), followed by the male (34%) and then by a second female (27%). The third female contributed very little at any stage. A picture of the male removing the egg was analyzed to estimate the egg maximum width, ruling out a possible runt egg. Motivations behind the behaviour of the male destroying the egg remain unclear.     

Seasonal declines in reproductive success of the common tern Sterna hirundo: timing or parental quality?

Reproductive success declines over the course of the breeding season in many bird species. Two categories of hypothesis have been evoked to explain this decline. The “timing” hypothesis suggests that seasonal declines in breeding success are attributable to the date of laying. The “parental quality” hypothesis suggests that seasonal declines result from the fact that young, inexperienced, or low quality birds breed later in the season. To evaluate the relative importance of timing and parental quality, egg exchanges and removals were used to manipulate hatching dates of common terns Sterna hirundo. Indices of quality, attendance, provisioning rates, and reproductive success of birds in three experimental groups (delayed hatch pairs, advanced hatch pairs, and pairs induced to relay) were compared to those of date‐matched controls. Pairs that hatched chicks early raised more chicks than pairs hatching chicks late in the season, regardless of initial laying date. This suggests that hatching chicks early is advantageous in itself. Our results, however, also support the parental quality hypothesis. There was a significant negative relationship between natural laying date and fledging success, independent of hatching date. Differences in chick growth and survival suggest that higher quality adults may be able to compensate for the disadvantages of late hatching dates and achieve similar reproductive success to that of pairs hatching chicks early. We found that pairs hatching chicks late in the season were subject to more incidents of kleptoparasitism than those hatching chicks early. This may be a proximate factor contributing to seasonal declines in reproductive success for common terns, although such a mechanism would not be likely in non‐colonial species. Failure to control for egg quality may have biased the results of some prior egg exchange experiments. Additionally, altered cost of incubation may be an unavoidable confounding factor in studies designed to manipulate timing of breeding.     

BREEDING BIOLOGY OF HOUSE MARTINS DELICHON URBICA IN RELATION TO AERIAL INSECT ABUNDANCE

The breeding biology of House Martins nesting in artificial boxes was examined in relation to aerial insect abundance and weather. The insect food supply was continuously monitored with a suction-trap at 40 ft above ground level. Aerial insect abundance started to rise from the winter level in April and reached a high stable state in June. Insects remained abundant until September; the subsequent decline continued at least until the end of October. The size of first clutches (mean size 3.87) was correlated with the abundance of aphids in spring, though not with temperature. It is suggested that this was an adaptive response because high food levels in May were associated with high levels in the main nesting period. Clutch-size declined through the season but was not matched by changes in food abundance. Second clutches (mean size 2–95) were most frequent following a high food level in June. No correlation was found between egg-weight and preceding food conditions, nor with season, clutch-size or order of laying within clutches. If food was scarce on the day of laying of the first egg, these clutches suffered a suspension of laying of subsequent eggs. Some of the young in these nests showed a reduced growth rate. The duration of incubation (mean 14.6 ± 1.1 days) was not apparently correlated with food abundance, however it must have been potentially extended by extreme shortages because incubation duties were neglected at such times. Infertility was probably the main cause of hatching failure (14.2%). A model was used to predict the date of onset of laying. It was based on the assumption that nestling tissue was produced with the same energetic efficiency as eggs. The model indicated that House Martins could collect enough food to lay eggs earlier than the observed date in most years. The high probability of food shortage before the laying period, however, appeared to discourage laying at this time. The observed onset of laying coincided with the appearance of aphids in the air, probably because they comprised an abundant and stable food source for egg formation. Breeding generally occurred in the period of highest food abundance. Pairs rearing only a single brood each year did so in July when food levels were highest. The growth of first brood nestlings was more closely associated with food levels than any other factor investigated, while in second broods, rainfall was found to have the greatest influence. The total variance in growth explained by environmental factors in the first broods (41%) was greater than in second broods (22%). This relative independence of second broods from adverse environments probably arose from more abundant food and the feeding of some of these broods by the young of first broods. An exceptionally late brood showed an extended nestling period indicating deteriorating conditions later in the breeding season. It is proposed that the spread of laying may reflect differences in the feeding efficiency of nesting House Martins, because the growth of two early first broods was more rapid at given food levels than two late broods. Laying patterns conformed to a distribution based on a progressive threshold mechanism. The more efficient pairs may be able to attain breeding condition and lay at low levels of food abundance, and hence breed earlier in the season. Subsequent layings are facilitated by the rise in food levels in early summer. Mortality within the nest was low (5.8%) and associated with food shortage. Nestling periods (mean 30.6 ± 2.3 days) were not shown to be correlated with food abundance; they were more dependent on brood-size. Recruitment into House Martin populations is thus widely influenced by food supply, particularly through an influence on clutch-size, the occurrence of second clutches and nestling mortality.