Juvenile helping in the moorhen,Gallinula chloropus

In a 3-year study of the moorhen, Gallinula chloropus, some chicks from first broods stayed on their natal territory once they had reached independence, and helped to rear their younger, second brood, siblings. Juvenile dispersal was constrained by habitat saturation, and first brood young were forced to stay on their natal territory. Juveniles that hatched early in the year were forced to stay longer, and helped more than those that hatched late. The total feeding rates to broods with and without juvenile helpers were the same, but parents with helpers reduced their feeding rates relative to parents without helpers. Pairs with helpers (=pairs attempting second broods) reared more chicks per nesting, attempt than pairs rearing chicks at the same time of year without helpers (=pairs attempting first clutch renests). This was true both for all clutches, and for hatched clutches only, even when controlling for parental quality, territory size and scasonal effects.     

Reproductive success of the Kittiwake Rissa tridactyla: the roles of clutch size, chick growth rates and parental quality

Kittiwake growth rates and breeding success are examined in relation to survival between fledging and breeding and to adult survival rates. High chick growth rates lead to increased survival after fledging. Broods of three (the maximum brood size) did not suffer lower fledging success than broods of two and clutches of three fledged appreciably more chicks per pair than did clutches of two or one. On average, the a- and b -chicks in broods of three grew at a faster rate and had a higher survival before breeding than those from smaller broods. Chicks from broods of two with experienced female parents grew at a faster rate than those of inexperienced female parents. Female parents which laid three egg clutches had a higher survival rate than those which laid clutches of two or one. We contend that three egg clutches were laid by higher quality individuals. We believe that clutch size indicates the condition of the Kittiwakes forming the pair. This condition probably has a genetical component, but is modified by environmental factors.     

Evolution of single-chick broods in the Swallow-tailed Gull Creagrus furcatus

Swallow-tailed Gulls lay single-egg clutches, and so raise single-chick broods. As they are pelagic seabirds, this small brood size is expected to relate to proximate food limitation owing to infrequent food deliveries. However, a previous brood doubling experiment detected an 82% increase in fledging success from experimentally doubled broods compared to controls. We repeated the brood doubling experiment, and found that none of 50 enlarged broods produced more than one independent offspring. Control and experimental parents produced fledglings of similar body size, which also had indistinguishable rates of fledging and subsequent survival and reproduction. A variety of parameters estimating survival and breeding costs of reproduction showed no treatment effect. Since two-chick broods yield dramatically higher fledging rates at some times, apparently without excess costs of reproduction, selection on brood size appears to favour a two-chick brood. However, selection may not favour a two-egg clutch if egg production is very costly. Additionally, our estimates of reproductive success do not incorporate the performance of experimental and control offspring as adults, which could differ, since growth of chicks differed slightly by treatment.     

The significance of clutch overlap in Great Tits Parus major

We studied the effects of manipulation of the size of first broods in the Great Tit Parus major on the size and breeding success of second clutches and its relation to the degree of clutch overlap. The rearing of first brood fledglings always overlapped with the laying of the second clutch and in most cases also with the incubation period of the latter. The degree of clutch overlap depended on the size of the first brood, being less when the first brood was large. Clutch overlap also increased with season. Mechanisms affecting the timing of laying of second clutches are discussed. A large first brood imposed reproductive costs. It affected the size of the second clutch by causing it to be delayed; second clutch size decreases with season. It affected the post-fledging survival of second brood young as, in this population, this decreases with fledging date. The breeding success of second clutches was, however, not affected by the size of the first brood, but instead by the weight of the female when rearing the first brood.     

Brood reduction in the Red-necked Grebe Podiceps grisegena

Brood reduction in Red-necked Grebes Podiceps grisegena breeding on fish ponds in south-eastern Poland occurred either through the desertion of the last-laid eggs after partial hatching of the clutch and/or the selective starvation of the smallest chicks. Abandonment of unhatched eggs was not influenced by the number of young already hatched or by the breeding date, but it was more likely in larger clutches and in families suffering chick starvation. Chicks from the largest broods had a higher probability of survival until fledging than those from single-chick broods. Larger chicks obtained food more successfully through better positioning during food delivery. In families that did not suffer brood reduction, chicks were better provisioned with food than in reduced broods. Although allocation of food among chicks in reduced broods was more skewed to the disadvantage of the younger siblings, dominant chicks obtained less food prior to brood reduction than dominant siblings in unreduced broods. Sibling aggression did not differ between unreduced and reduced broods before death of the weakest chicks. Post-laying adjustment of the number of offspring to prevailing feeding conditions occurred at two stages: by parental manipulation of the number of hatched eggs at the time when parents and chicks leave the nest and by competition between chicks. It is suggested that late egg desertion may be an adaptive mechanism of brood-size adjustment, when elimination of the weakest chicks through sibling competition is not very efficient.     

Nestling growth and mortality of Pied Flycatchers Ficedula hypoleuca in relation to weather and breeding effort

The growth pattern and mortality of young Pied Flycatchers Ficedula hypoleuca were studied to focus on the mechanisms and constraints behind the widely studied optimization of clutch size. The clutch sizes were modified, and the growth and survival of chicks from different clutch sizes were monitored along with the prevailing weather during the nestling period to detect the effect of weather on reproductive success. The weather conditions during the feeding period of the nestlings varied within a season as well as between breeding seasons. The prevailing weather markedly affected both the growth rate and the survival of chicks, yet, the effects of weather on growth were not greater in enlarged clutches. The impact of adverse weather was more pronounced in the later phases of nesting, when the food demand of a brood was highest. Brood reduction and total nest losses were more likely during extensive rainfall during the nestling phase and also in the enlarged clutches. Thus, weather is a very important determinant of reproductive success in this species. Weather conditions during the breeding season are unpredictable, however, and therefore brood reduction through sibling competition is a mechanism whereby brood size can be adjusted to the level the parents can rear under the prevailing conditions.     

THE SOCIABLE WEAVER,PART 4: PREDATORS,PARASITES AND SYMBIONTS

Maclean, G. L. 1973. The Sociable Weaver, Part 4: Predators, parasites and symbionts. Ostrich 44: 241–253.

The main nest predator of the Sociable Weaver in the Kalahari sandveld is the Cape Cobra Nala nivea. This snake causes great losses of eggs and chicks; one cobra may eat the contents of an entire nest mass at one feed. Another nest predator which causes smaller losses of eggs and chicks but great destruction to the nest masses is the Honey Badger Mellivora capensis. These are the only two nest predators on the Sociable Weaver in the study area. Predators on adult Sociable Weavers include several birds of prey and some small carnivorous mammals.

Adult Sociable Weavers have few ectoparasites and hardly any Mallophaga. A common ectoparasite on the legs of chicks is a blood-sucking Dermestes larva, which appears not to be harmful. The only endoparasite found was the nematode, Diplotriaena ozouxi, which infected the abdominal air sacs of the adults.

The nest material of the Sociable Weavers' communal nest masses was inhabited by a wealth of invertebrate animals and a few harmless reptiles such as skinks and geckos.

Some of the chambers in a Sociable Weaver nest mass may be taken over by other species of birds. Most of these, such as Redheaded Finches Amadina erythrocephala, use the chambers for breeding purposes only, but the most important avian symbiont, the Pygmy Falcon Polihierax semitorquatus, is a permanent resident in the chambers. The presence of Pygmy Falcons is resented by the weavers but the falcons may help to keep snakes away from the nest mass. Adult Sociable Weavers are not normally preyed on by Pygmy Falcons, although the falcons may occasionally take young weavers in the nest chambers.

The tops of the nest masses may be used as nest sites by the Giant Eagle Owl Bubo lacteus. Barn Owls Tyto alba may use cavities in the superstructure of nest masses for roosting in. Neither of these owls appeared to prey on the weavers.     

BREEDING BIOLOGY OF THE EDIBLE-NEST SWIFTLET AERODRAMUS FUCIPHAGUS

A colony of the Edible-nest Swiftlet Aerodramus fuciphagus nesting in a Chinese shophouse in Penang, Peninsular Malaysia, was studied for seven months. Birds bred throughout, but laying was concentrated in the period October to February. Incubation and fledging periods were 23 ± 3 days and 43 + 6 days, respectively. Most young hatched during the dry season. The normal clutch size was two, with many birds laying second clutches (75%) and some third clutches (15-4%) using the same nest. Clutch size showed no variation with clutch order or month. The overall hatching success was 69-0 %, most losses caused by eggs falling from or with the nest. Fledging success was similar for broods of two (625%) and broods of one (59-4%), so that the former were twice as productive. The main loss of chicks occurred when they fell from the nest and were eaten by mammalian predators. There was a tendency for breeding success to decline with successive clutches, but not with month, being highest in November (53-6 %) and February (55-6 %). The growth rate of single chicks and successful broods of two was similar, except that the second chick was more prone to fluctuations in weight. Some second chicks showed evidence of starvation before falling from the nest. Diet was examined by analysis of foodballs regurgitated by mist-netted adults. Foodballs weighed 0–13-1 08 g and, on average, contained over 500 prey items. The main arthropods caught were Hymenoptera (40-8%), Ephemeroptera (26-4%), Homoptera (15-4%) and Diptera (7-7%). Flying ants and mayflies contributed most by weight, although figwasps and mayflies were the most numerous prey items. Only in December did the percentage of moulting mist-netted adult swiftlets fall below 70 %. It appeared that many birds were moulting and breeding simultaneously, although 21% of birds had primaries moulting in two places, suggesting arrested moult. Diurnal activity showed a normal high dawn exodus and dusk inflow of birds, although there was evidence of an increase of birds flying out prior to the dusk inflow. Nest harvesting was continued throughout the study, but most of the marked nests under study were left undisturbed.     

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.     

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.     

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.     

The breeding biology of the Pacific Swallow Hirundo tahitica in Malaysia

The breeding biology of the Pacific Swallow was studied in Malaysia. Time-budgets and preliminary energy-budgets were calculated for different stages of the breeding cycle. Breeding was seasonal, the first eggs were laid in the first week of March from 1978 to 1982. The nest was a mud cup built under an overhang, often on man-made structures over water. The mean clutch size was 2.98 eggs. So more than two successful clutches were ever raised, although up to four sets of eggs can be laid if clutches are lost. The incubation period was 16.2 days and the nestling period 19–21 days. Nestling weights showed the typical hirundine recession before fledging. The mean brood size was 2.32 and there was no evidence of undernourishment in larger broods. Nesting success was lower than for temperate hirundines but high for a tropical passerine. Incubation duties were carried out by the female alone but both parents fed the brood. The male never delivered more than 44% of the total feeds. Daily energy expenditure varied from 3.44–4.9×BMR depending upon the stage of the breeding cycle.     

Age-dependent reproductive costs and the role of breeding skills in the Collared flycatcher

This study addressed whether there are any age‐related differences in reproductive costs. Of especial interest was whether young individuals increased their reproductive effort, and thereby their reproductive cost, as much as older birds when brood size was enlarged. To address these questions, a brood‐size manipulation experiment with reciprocal cross‐fostering of nestlings of young and middle‐aged female Collared flycatchers, Ficedula albicollis, was performed on the Swedish island of Gotland. Nestlings’ body mass, tarsus length and survival were recorded to estimate the parental ability and parental effort of the experimental female birds. Female survival and clutch size were recorded in the following years to estimate reproductive costs. We found that middle‐aged female flycatchers coped better with enlarged broods than younger females or invested more in reproduction. In the following year, young female birds that had raised enlarged broods laid smaller clutches than the females from all the other experimental groups. This result shows that the young female birds pay higher reproductive costs than the middle‐aged females. Both young and middle‐aged female flycatchers seemed to increase their reproductive effort when brood size was increased. However, such an increase resulted in higher reproductive costs for the young females. The difference in reproductive costs between birds of different ages is most likely a result of insufficient breeding skills of the young individuals.     

Fitness cost of incubation in great tits (Parus major) is related to clutch size

Life-history theory predicts that parents produce the number of offspring that maximizes their fitness. In birds, natural selection on parental decisions regarding clutch size may act during egg laying, incubation or nestling phase. To study the fitness consequences of clutch size during the incubation phase, we manipulated the clutch sizes during this phase only in three breeding seasons and measured the fitness consequences on the short and the long term. Clutch enlargement did not affect the offspring fitness of the manipulated first clutches, but fledging probability of the subsequent clutch in the same season was reduced. Parents incubating enlarged first clutches provided adequate care for the offspring of their first clutches during the nestling phase, but paid the price when caring for the offspring of their second clutch. Parents that incubated enlarged first clutches had lower local survival in the 2 years when the population had a relatively high production of second clutches, but not in the third year when there was a very low production of second clutches. During these 2 years, the costs of incubation were strong enough to change positive selection, as established by brood size manipulations in this study population, into stabilizing selection through the negative effect of incubation on parental fitness.     

The feeding and breeding ecology of Barn Owls Tyto alba in Peninsular Malaysia

Barn Owls have only recently colonized Peninsular Malaysia, nesting in the roof spaces of houses in oil palm estates and feeding on the rats which inhabit these plantations. Pellet analysis showed that the prey spectrum was confined almost entirely to three species of the genus Rattus which are the major pests of oil palm. There was no annual variation in diet. Breeding showed a broad seasonality but occurred in all months of the year. Mean clutch and brood sizes of 6.6 and 4.6 respectively were recorded, most pairs producing two broods a year although on two occasions three were raised. Overall hatching success was 69.0% with first clutches more successful (79.9%) than second (57.3%). First broods fledged 86.1% and second broods 69.1% of young fledged. Comparison of growth rates of different sized broods suggested that there is a physiological maximum at which all broods proceed irrespective of brood size. The behaviour al changes needed in hunting techniques when colonizing dense plantations rather than the more usual open habitat of Barn Owls is discussed. The breeding strategy seems to be one of producing large clutches and broods, and frequent breeding attempts in a habitat with a high potential carrying capacity.     

Strong evidence for selection for larger brood size in a great tit population

We measured the selection pressure on brood size in a recentlyestablished population of great tits (Parus major L.) in thenorthern Netherlands by manipulating brood size in three years(1995: n = 51, 1997: n = 66, 1998: n = 51), and we estimatedfitness consequences in terms of local survival of both offspringand parents. Enlarged broods had highest fitness; the offspringfitness component was positively affected by manipulation andthe parental fitness component was unaffected. Parental survivaland the probability that parents produced a second clutch werenot affected by the treatment. However, parents that had raisedenlarged broods produced their second clutch later in the season.Clutch size, brood size, and laying date of birds recapturedin the next breeding season were largely independent of thetreatment. We conclude that there is strong evidence for selectionfor larger brood size and reject the individual optimizationhypothesis for this population because the number of young inthe nest predicts fitness independently of the manipulationhistory.     

Sociable Weaver biometrics and primary moult

The biometric and primary moult data housed at the South African Bird Ringing Unit (SAFRING) were analysed for the Sociable Weaver Philetairus socius. The average body mass and wing length was 27.9g (SD = 2.2) and 74.1mm (SD = 2.5), respectively. Variation in these parameters is not clearly correlated with region, season or climate, other than a negative correlation of body mass with average annual water deficiency. Body mass of Sociable Weavers near Kimberley showed a longterm decrease of 2.9g, probably due to stabilising selection on mass. Primary moult duration varied from 152 days to 169 days and started between 26 January and 31 December in two populations (socius and South African eremnus respectively). Individual primaries moulted mainly one at a time, each taking 20–28 days to grow fully. Prolonged moult duration in this species is probably an adaptation to reduce energy expenditure, and to grow more durable feathers due to abrasion in entering the nest. The lack of clear patterns of geographical variation in biometrics indicates that the contiguous populations of Sociable Weaver should belong to the nominate species. The biometric and primary moult data housed at the South African Bird Ringing Unit (SAFRING) were analysed for the Sociable Weaver Philetairus socius. The average body mass and wing length was 27.9g (SD = 2.2) and 74.1mm (SD = 2.5), respectively. Variation in these parameters is not clearly correlated with region, season or climate, other than a negative correlation of body mass with average annual water deficiency. Body mass of Sociable Weavers near Kimberley showed a longterm decrease of 2.9g, probably due to stabilising selection on mass. Primary moult duration varied from 152 days to 169 days and started between 26 January and 31 December in two populations (socius and South African eremnus respectively). Individual primaries moulted mainly one at a time, each taking 20–28 days to grow fully. Prolonged moult duration in this species is probably an adaptation to reduce energy expenditure, and to grow more durable feathers due to abrasion in entering the nest. The lack of clear patterns of geographical variation in biometrics indicates that the contiguous populations of Sociable Weaver should belong to the nominate species.     

ASPECTS OF THE BREEDING BIOLOGY OF THE FIERYNECKED NIGHTJAR

Jackson H. D. 1985. Aspects of the breeding biology of the Fierynecked Nightjar. Ostrich 56: 263–276.

A marked population of nightjars in Zimbabwe was studied intensively for four breeding seasons. This paper covers certain aspects of the breeding biology of the Fierynecked Nightjar Caprimulgus pectoralis. The male shows strong site fidelity during the breeding season (September to December), singing, feeding and breeding within an area of about 5,8 ha. There is some evidence of site defence by the male. The female shows strong mate fidelity, resulting in a pair bond for life. Egg laying starts with full moon in September and is further stimulated by the next two full moon periods. The eggs are laid directly on dense leaf litter at a site overhung by foliage. The normal clutch is two eggs (12S % are one egg) laid on successive days during the afternoon. Incubation starts with the first egg and is by the male at night and the female by day. The incubation period is 18 days. The birds respond to undue disturbance by deserting the eggs and laying a replacement clutch. The chicks usually hatch on successive afternoons; they are mobile on the first day and will move to a parent if called. Both parents feed and brood the young during twilight and moonlight; the male broods them on dark nights and the female does so by day. The species is double-brooded when time permits, the female laying again once the first brood has reached independence; she may even lay a third clutch if the second one comes to grief. There is no evidence of adults transporting eggs or young.     

Interval between clutches, fitness, and climate change

Timing of optimal reproduction can be affected by the presenceof multiple broods, with multi-brooded species breeding earlier(and later) than the optimal timing of breeding as comparedwith single-brooded species that only need to optimize the timingof a single brood. Approximately two-thirds of barn swallowsHirundo rustica produce 2 broods per year, and I tested whetherthe constraints on timing of reproduction were affected by climatechange because climatic amelioration would allow both an earlierstart and a later termination of reproduction. The durationof the interval between first and second clutches and the variancein the duration increased during 1971–2005 when temperatureduring spring, but not summer, increased rapidly. Interclutchinterval was shorter when mean date of breeding was late andalso among late-breeding individuals during individual years.When clutch size and brood size of the first clutch were large,interval until the second brood increased. Pairs with a longinterval produced more fledglings than pairs with a short interval.Pairs with first broods with strong mean T-cell–mediatedimmune responses took shorter time to start their second clutch,whereas mean body mass or tarsus length of first broods werenot significantly related to interclutch interval. Interclutchinterval increased with the size of a secondary sexual character,the length of the outermost tail feathers of adult male barnswallows, but not with tail length of females, or with sizeof several other phenotypic characters in either sex. Thesefindings are consistent with the hypothesis that the durationof the interclutch interval is determined by a combination ofenvironmental conditions, reproductive effort, and sexual selection.     

Annual cycle of the Helmeted Honeyeater Lichenostomus melanops cassidix, a sedentary inhabitant of a predictable environment

The annual cycle of breeding, moult and weight variation in the Helmeted Honeyeater Lichenostomus melanops cassidix , a sedentary bird of temperate southeast Australia, is documented. Breeding and moult were sequential unimodal annual events, whose timing was highly consistent between years. However, overlap of breeding and moult was frequent, and some individuals even commenced primary moult before laying their final clutch. The timing of the post-juvenile moult was coincident with that of adults. Early-hatched young moulted within a few months of hatching, but late-hatched young deferred moult for a year. Helmeted Honeyeaters were heaviest in autumn and early winter, and lightest in spring and early summer, a cycle most consistent with the redirection of all available resources to reproduction. The long breeding season (seven-and-a-half months) of the Helmeted Honeyeater, extensive overlap of breeding and moult, and other life-history attributes including small clutch size, are more consistent with the described bio-rhythmic patterns for birds in the humid tropics than the temperate zone. However, the Helmeted Honeyeater has a fairly rapid primary moult rate, unusual amongst species that overlap moult and breeding. This combination of attributes reflects the stable, somewhat seasonal environment occupied and the resource monopoly established by this tightly territorial subspecies. We speculate that extension of the breeding season, by overlapping breeding and moult, is one of the few options available to vary life-history strategies amongst 'old-endemic' Australian birds of the temperate zone.