Effects of clutch size and egg-laying order on the breeding success in the Little Tern Sterna albifrons on the Nakdong Estuary, Republic of Korea

Survivorship of Little Tern Sterna albifrons eggs and chicks was followed on an islet in the Nakdong Estuary, Republic of Korea, in 1995 and 1996. Mean egg size and incubation period were significantly different between the 2 years. The maximum clutch size was three eggs, and the second egg in the clutch often hatched earlier than the first, while most of the third eggs hatched last. In 1996, when the fate of 249 eggs from 106 nests was followed for 40 days, hatching success, fledging success and breeding success were 77%, 40% and 31%, respectively. High mortality occurred in the early chick stage, mostly because of rain and predation by Weasels Mustela sibirica. The breeding success per egg was 14% in one-egg clutches, 28% in two-egg clutches and 34% in three-egg clutches. This difference was mainly attributed to the lower hatching success in the smaller clutches. In three-egg clutches, the third egg showed significantly lower breeding success than siblings. The main foods of the Little Tern were Tridentiger obscurus, Engraulis japonicus, Hyporhamphus intermedius, Acanthogobius flavimanus (all fish), Palaemon sp. and Crangon affinis (shrimps). The feeding frequency was, apparently, not affected by time of day and age of chicks but was probably influenced by weather conditions. Newly hatched chicks failed to eat 25% of the prey brought to them, although this decreased with the age of the chicks.     

Population dynamics and reproduction of Northern Lapwings Vanellus vanellus in a meadow restoration area in central Sweden

To investigate the effectiveness of a meadow restoration programme, we studied the relationships between population changes and environmental changes, return rates and hatching success in a population of Northern Lapwings Vanellus vanellus. The study was performed on mixed farmland (59 km²) in central Sweden, an area that underwent extensive meadow restoration between 1985 and 1994. The study included more than 2600 nests, supplemented with observations of 127 individually colour-ringed Northern Lapwings. The breeding population varied (2.7–5.3 pairs/km²), but showed no significant trend with time. The population increased in years with high spring flooding levels. Population size was unrelated to demographic factors (e.g. hatching success the previous year (14–50%), and return rate). Lapwings moved considerably between years and their nest site fidelity was unrelated to previous hatching success or other factors, suggesting that changes in habitat quality and migration between populations were important in regulating population size. Recent extensive meadow restoration did not seem to aid the Northern Lapwing population; birds continued nesting on tillage even though most nests were destroyed by farming activities. A relatively high relaying frequency improved hatching success, which was still lowest in the most preferred habitats (spring sown crops, total hatching success c. 30%). The few Northern Lapwings breeding in the least preferred habitats (meadows and cultivated grassland) had a better hatching success (> 70% total hatching success), suggesting that habitat selection was not determined by hatching success. From these results, recommendations are made for the management of Northern Lapwing (and associated species) populations on farmland.     

Long-term trends in the abundance of breeding Lapwing Vanellus vanellus in relation to land-use change on upland farmland in southern Scotland

Capsule A long-term decline on this habitat is shown to be associated with the intensification of agricultural management, particularly the occurrence of field drainage.

Aims To assess long-term trends in the number of breeding Lapwing and determine the relationship between these trends and changes in agricultural management on an upland study area.

Methods Breeding Lapwing were counted along two road transects in nine years between 1980 and 2002, and on one extensive plot in 1980, 1990 and 2000. Counts along the road transects were made from a vehicle and the fields used for nesting were recorded. Changes in field management along the transects were monitored annually between 1980 and 1990, and habitat composition assessed in 1980, 1985, 1990 and 2000.

Results During the first 20 years of study the number of breeding Lapwing declined substantially on all three count areas and by 77% overall, with further declines on both transects in 2002. The area of unimproved grassland and arable on these transects also declined substantially due to conversion to improved grass. Fields that comprised either unimproved grassland or arable were most likely to hold nesting Lapwing, while the chance of a field losing its nesting Lapwing was positively associated with the occurrence of drainage. Drainage and conversion to improved grass were closely linked.

Conclusions Agricultural intensification is a probable cause of decline in the number of breeding Lapwing in upland areas. Such declines may have been widespread in upland areas following increased agricultural intensification in recent decades.     

THE BIRDS OF LANGEBAAN LAGOON

Data are presented on breeding success of Red Bishops (Euplectes orix) collected over four breeding seasons at a colony in the Addo Elephant National Park, Eastern Cape, South Africa. Overall hatching and fledging success were 53.8% and 26.0% of all eggs laid, respectively, and the overall mean number of fledglings per breeding attempt was 0.77. Hatching and fledging success varied significantly among seasons, with both clutch and brood losses due to predation being the main reason for the observed differences. Hatching success also differed significantly among clutch sizes, being highest for four-egg clutches (63.2%), intermediate for three-egg clutches (55.5%) and lowest for two-egg clutches and five-egg clutches (33.2% and 34.3%, respectively). However, fledging success was not significantly different among clutch sizes. The mean number of fledglings per breeding attempt was 0.44 for two-egg clutches, 0.80 for three-egg clutches, 1.10 for four-egg clutches, and 0.57 for five-egg clutches. The height of accepted nests (i.e.nests in which at least one egg was laid) was significantly lower than the height of nests not accepted. In addition, accepted nests in which eggs hatched and young fledged were significantly lower than accepted nests in which no eggs hatched and no young fledged. These overall effects of nest height on nest acceptance and hatching and fledging success were, however, due only to nests built above water, since no such effects were found when nests built above ground (i.e.on dry land) were analysed separately. I detected no effect of nest coverage on the probability of a nest being accepted, nor was there any effect of nest coverage on hatching or fledging success. Nests above water were significantly more likely to be accepted than nests above ground; however, hatching and fledging success of nests that were accepted did not differ significantly between nests built above water and those built above ground.     

Costs of large communal clutches for male and female Greater Rheas Rhea americana

The breeding system of the Greater Rhea Rhea americana is almost unique among birds as it combines harem polygyny and sequential polyandry, with communal egg-laying and uniparental male care. In this species, large communal clutches (more than 30 eggs) are rare and have a lower hatching success than smaller clutches. Here we analyse the proximate causes of hatching failures and the costs of large communal clutches (and therefore the costs of extensive polygyny) for males and females. We evaluated if length of the nesting period, egg viability, egg losses during incubation and male parental activity at the nest were affected by clutch size. We also evaluated if chicks hatched from large clutches have a lower survival during the first 2 months after hatching. Large clutches had longer nesting period and lower hatching success, mainly as a result of bacterial contamination of the eggs and increased hatching asynchrony. In addition, large clutches tended to lose more eggs as a result of accidental breakage or predation. Male activity at the nest and chick survival were not related to clutch size. Low hatching success, nest predation risk and energetic costs associated with large clutches penalize females that join large harems and males that accept additional eggs into the nest.     

Rates and consequences of relaying in little auks Alle alle breeding in the High Arctic an experimental study with egg removal

Most seabirds have a small clutch size. Thus, replacement of a clutch after loss can make important contributions to an individual’s lifetime reproductive success. However, in the condition of short polar summer, relaying propensity may be time‐constrained. In this study, we investigated rates and consequences of relaying in a small High Arctic seabird, the little auk Alle alle. We performed an experiment in which we removed the single egg from 20 nests of early‐laying breeders. We measured relaying rates, and compared chick body mass and breeding success between the experimental and control nests. Despite the narrow window of the Arctic summer and the closely synchronized breeding, 75% of females produced a replacement egg just 2.7% smaller in volume than the first egg. This indicates that in little auks, the demographic effects of disruptions to breeding attempts (by predators, adverse weather or human activity) may be mitigated to some extent by replacement clutches. However, peak body mass and fledging body mass were lower in the experimental than the control chicks. This effect was rather a consequence of late hatching – chicks from replacement clutches followed seasonal decline in peak body mass and fledging mass. Finally, breeding success and chick survival up to 20 d in the experimental nests were respectively 34 and 37% lower than in the control nests. Thus, the quality and post‐fledging survival of chicks from the replacement clutches were probably lower compared to the chicks hatched from the first‐laid eggs.     

The use of predator-exclusion fencing as a management tool improves the breeding success of waders on lowland wet grassland

Waders breeding on lowland wet grassland have undergone dramatic declines across Europe in recent decades. Few species now achieve the levels of breeding success required for population stability and recovery, with predation from large mammals acting as a key compounding factor limiting nest survival and productivity. Predator management through lethal control is often controversial, yet alternative non-lethal methods are little tested in the context of grassland breeding waders. Excluding predators through the use of electric fences has led to improvements in nest and chick survival in other habitats. To test the applicability of this method to lowland wet grassland we constructed predator-exclusion fences on sites across the UK and, with Lapwing Vanellus vanellus as a study species, used historical and contemporary data to test whether excluding large mammalian predators leads to an increase in wader nest survival and productivity, and whether effects differ between fence designs. Lapwing nest survival was significantly higher in the presence of any type of predator-exclusion fence, with significantly fewer nests predated each day. Overall productivity also improved, with significantly higher numbers of chicks fledged per pair in years when fences were operational. Different designs and methods of powering fences resulted in different levels of success, with combination design fences and those powered by mains electricity performing best. Excluding large mammalian predators from areas of lowland wet grassland with predator-exclusion fencing successfully improves Lapwing nest survival and productivity, allowing breeding success to exceed the levels required for population recovery. Other wader species breeding in the same habitat are also likely to benefit from the increased protection from predation provided by fences. Predator-exclusion fencing is therefore an effective management tool for protecting restricted and declining populations of breeding waders on lowland wet grassland.     

Seasonal variation in reproductive measures of tropical Roseate Terns Sterna dougallih previously undescribed breeding patterns in a seabird

Seasonal variation in egg-laying, egg size, hatching success, hatchling mass, fledging success and chick growth of Roseate Terms Sterna dougallii breeding on Aride Island (Seychelles), Indian Ocean, were studied in 1997 and 1998. I investigated to what extent two patterns, common in a range of species, were followed by tropical Roseate Terns: (a) seasonal decrease in clutch size, egg size and breeding success and (b) an increase in breeding success with increasing egg weight. In 1997 (a poor year), the earliest nesting birds laid significantly smaller eggs, and chicks were lighter at hatching than those of peak nesting birds. The mean clutch size, of 1.04 eggs, showed no seasonal variation and no 'b'-eggs hatched. In 1998 (a good year) the earliest nesting birds laid eggs of similar size and their chicks were of similar weight to those of peak nesting birds. Mean clutch size, of 1.25 eggs, increased significantly through the season and about 60% of the 'b'-eggs hatched. In 1997, hatching success was 57% whereas in 1998 it was 80%. In both years, breeding success declined significantly through the season. The fact that the earliest breeding birds laid smaller eggs in a poor year and smaller clutches in a good year is in marked contrast to a range of other species, and to temperate-nesting Roseate Terns. Egg volume explained about half of the variance in hatchling mass in both years, but only 15% of the variation in linear growth rate. Hatching date was the only variable with a significant effect on fledging success. Roseate Terns on Aride seemed to sacrifice egg size and clutch size for earliness of laying. Presumably it is a strategy of older birds to lay as early as possible and may be regarded as a response of tropical Roseate Terns to breeding under relatively poor, and seasonally declining, food conditions.     

Breeding success of the Eurasian eagle owl (<Emphasis Type="Italic">Bubo bubo</Emphasis>) and rodent population dynamics

The dependence of the breeding success of the eagle owl on the population dynamics of rodents, which are a staple of its diet in Mordovia, was traced. At peak numbers of rodents, large clutches and a high survival rate of fledglings were observed in the breeding pairs; after a year of depression of rodent populations, in the following year, the pairs do not nest at all or their breeding success is reduced to a minimum due to the death of the clutches. In 52% of the cases, the nesting pairs laid three eggs; in 31%, two eggs; in 9%, four eggs; in 4%, one egg or five eggs. The average clutch size was 2.78 ± 0.17. The average number of chicks grown from a successful nest was 2.41 ± 0.27. The nests in Mordovia were located at a distance of 1.1–3.7 km from residential areas.     

Breeding success of a colony of Boat-billed Herons Cochlearius cochlearius (Ciconiiformes: Ardeidae) in pasturelands of Costa Rica

The breeding success of a double-brooding colony of Boat-billed Herons Cochlearius cochlearius was studied in pasturelands of Costa Rica. Mean clutch size in the first clutches (2.9 eggs/nest) was higher than in second and repeat clutches (2.3 eggs/nest). Breeding success was similar in the first attempt and second attempts (20.7% and 21.7%, respectively). In both attempts earlier nests enjoyed a higher breeding success. Starvation of the youngest chicks within the nest and destruction of nests by bad weather conditions were the main factors related to nestling death. No effects of human activity on the reproduction of the breeding colony were observed.     

Exclusion of ground predators improves Northern Lapwing Vanellus vanellus chick survival

Many farmland‐breeding wader species have declined across Europe, probably due to reductions in reproductive output caused by high nest losses as a result of agriculture or predation, or low chick survival between hatching and fledging. Most studies have focused on nest failures, and the factors affecting post‐hatching survival of chicks are poorly known. In an experimental approach, we fenced parts of the arable foraging areas of Northern Lapwing Vanellus vanellus families to quantify chick survival simultaneously in the presence and absence of ground predators. Lapwing chicks were radiotagged to estimate survival probabilities by daily locations, applying multistate capture–recapture models. During the night, chick survival was considerably lower outside fenced plots than within. During the day, chick survival was higher than at night and did not differ between protected and unprotected plots. This suggests that nocturnal ground predators such as Red Foxes Vulpes vulpes were responsible for a significant proportion of chick mortality. Cumulative survival probability from hatching to fledging was 0.24 in chicks within fenced plots, but virtually zero in chicks outside fenced plots. In farmland, temporary electric fences can be effective in minimizing the impact of ground predators and offer a promising short‐term method to increase fledging success of precocial birds.     

ASPECTS OF McCORMICK SKUA BREEDING BIOLOGY

The breeding behaviour of the McCormick Skua was studied at Cape Royds, Antarctica, for three summers. These observations followed the work of Young (1963a, b). Records of banded birds suggest that at the age of five to six years birds return to breed in the area where they were reared. Breeding pairs retain the same territory and mate over consecutive summers. The male will retain a territory and remate if the female does not return, while there is some evidence to suggest that if the male is absent the female will leave the territory and remate elsewhere. Unfavourable weather frequently forced incubating birds to desert their eggs and as a result some pairs laid three clutches (at the most two clutches of two eggs and a clutch of one). The death of sibling chicks from fighting was examined by comparing the early growth of 11 successful pairs of chicks with that of 12 pairs in which one of the siblings died as a result of fighting. From a comparison of weights at hatching and the interval between the hatching of members of chick pairs, it appears that the difference in weight between twins and the gain in weight by one chick in the first few days is critical for the survival of the twins. The breeding success, calculated from the total eggs laid and the total number of chicks fledging, was 40%, 42% and 17% for the three respective summers. Storms (periods of wind with snow) were the main factor influencing breeding success.     

Factors influencing the nesting success of Lapwings Vanellus vanellus and behaviour of Red Fox Vulpes vulpes in Lapwing nesting sites

Capsule Lapwing nest predation was negatively correlated to nest density, while Lapwing alarm duration in response to foxes was positively correlated with the number of Lapwing broods present.

Aims To identify factors affecting Lapwing nest predation and Red Fox search effort.

Methods Lapwing nest success was monitored at four sites in 1996, seven sites in 1997 and six sites in 1998. In 1997 we mapped the position of all Lapwing nests in order to determine distances between nests, and the proximity of linear features and potential avian predator perches to each nest. From April to June 1998 we carried out 199 hours of nocturnal observations at six Lapwing nesting sites using night vision equipment.

Results The risk of nest predation was significantly higher for more isolated nests. Nocturnal observations showed that of all the nocturnal predators, foxes were the most active at Lapwing nesting sites. However, fox search effort in Lapwing colonies was relatively low, averaging 57 s/ha per visit. Foxes spent significantly longer foraging near breeding Lapwings (measured as duration of alarm calls) when more broods were present. Fox search effort (s/ha per hour of observation) tended to be greater in areas of high waterbird density.

Conclusion The lack of positive density-dependent nest predation, the relatively low search effort of foxes near Lapwing nesting sites and the high nest success sometimes achieved in areas with foxes all suggest that Lapwing nest predation by foxes is ‘incidental’. Lapwing chicks are probably more vulnerable to predation by foxes than clutches.     

Supplementary feeding increases Common Buzzard Buteo buteo productivity but only in poor‐quality habitat

Temporal heterogeneity in the effects of food supply during the breeding season on the productivity of the Common Buzzard Buteo buteo was investigated in a supplementary feeding experiment. Pairs were fed artificially (1) before egg‐laying, (2) after chicks hatched and (3) continuously throughout the season, and compared with (4) unfed controls. Pairs fed before egg‐laying had marginally larger clutches than those not fed, but lay date, egg volume and weight, brood size and hatching success were unaffected. Territorial quality had far greater effects, with pairs nesting in low‐quality habitats (bog, scrub and semi‐natural grassland) laying later and having lower hatching success, smaller broods and fewer fledglings than those in more productive agricultural landscapes. Supplementary feeding after egg hatching neutralized the negative effect of poor habitat, resulting in fed birds having significantly more fledglings. This study emphasizes the importance of food availability when provisioning chicks in suboptimal habitats and has implications for the success of diversionary feeding in reducing game losses to Buzzards.     

Intraclutch egg-size variation and offspring survival in the Kentish Plover Charadrius alexandrinus

We studied the consequences of intraclutch egg-size variation in Kentish Plovers Charadrius alexandrinus in southern Spain to test the hypothesis that females allocate resources preferentially to eggs with the greatest survival potential. Second eggs were larger and had a greater hatching success than the other eggs in a clutch. However, we did not find unequivocal support for the differential allocation hypothesis, because egg failures according to laying order were not due to the causes predicted by the hypothesis, and the change in egg-size with laying order was not consistent between successive clutches of renesting females. This suggests that nutrient availability during egg formation could act as a proximate factor affecting egg-size independently of laying order. Larger eggs produced heavier chicks and, within broods, heavier chicks were most likely to be recruited into the breeding population. By laying clutches with a larger mean egg volume in replacement clutches, females took longer to lay again after failure of the first clutch. Thus, opposing selective forces may act to increase and decrease egg volume. Although a trade-off between egg-size and clutch-size has not been found in precocial birds, our results suggest that there may be a trade-off between egg-size and fecundity, since an increase in the size of eggs within clutches may limit the time remaining within the season to lay second or replacement clutches.     

Effect of brood size and hatching sequence on prefledging mortality of Sandwich terns: why lay two eggs?

The mortality of Sandwich tern Sterna sandvicensis chicks held in enclosures was studied in colonies on Griend, in the Dutch Wadden Sea, from 1992 to 1999, and on Hirsholm, in the Danish Kattegat, in 1997. Survival of chicks until fledging was 73% for chicks hatching from first-laid eggs or single-egg clutches and 59–64% for partially hatched two-egg clutches, whereas 6% of second hatchlings survived until fledging. Less than 2% of all two-chick broods actually fledged two chicks. Because 18% of the two-egg clutches only hatched one egg, 7% of fledglings of two-egg clutches originated from a second-laid egg. In nests where both eggs hatched, the number of chicks was usually reduced soon after hatching. Within five days of hatching more than 50% of the second hatchlings died of starvation or were preyed upon. It seems that overproduction commonly occurs in Sandwich terns and that investment in a surplus egg mainly serves as an insurance mechanism. On Griend and Hirsholm, chick productivity of two-egg clutches was somewhat higher than for one-egg clutches. Undernourishment was an important cause of death, either directly by starvation or by selective predation of chicks in poor condition. This, in combination with earlier, studies suggests that Sandwich tern parents on Griend are exposed to severe food stress.     

Age‐dependent dispersal and habitat choice in black‐tailed godwits Limosa limosa limosa across a mosaic of traditional and modern grassland habitats

Whether to disperse, and where to, are two of the most prominent decisions in an individual's life, with major consequences for reproductive success. We studied natal and breeding dispersal in the monogamous black‐tailed godwit Limosa limosa limosa in the Netherlands, where they breed in agricultural grasslands. The majority of these grasslands recently changed from wet herb‐rich meadows into well‐drained grassland monocultures, on which godwits have a lower reproductive success. Here we examine habitat selection with a multistate mark–recapture analysis. Habitat transition probabilities between meadows and monocultures were estimated on the basis of 1810 marked chicks and 531 adults during seven years in a 8500 ha study area. Young and adult godwits may differ in habitat selection because: 1) adults may have gained experience from previous nest success where to settle, 2) younger individuals may find it harder to compete for the best territories. Both young and adults moved at a higher rate from the predominant monocultures to meadows than the other way around, thus actively selecting the habitat with better quality. However, dispersal distance of adults was not affected by previous nest success. The average dispersal distance from place of birth of godwits breeding for the first time was ten times larger than that of adult godwits. That godwits breeding in their second calendar year arrived and laid at similar dates and were equally able to select territories in areas with high breeding densities, suggests that young birds were not competitively inferior to adults. Although on monocultures reproduction is insufficient to maintain constant populations, birds sometimes moved from meadows to monocultures. This explains why even after 30 years of land‐use intensification, godwits still breed in low‐quality habitat. The adjustment to changing habitat conditions at the population level appears to be a slow process.     

Maternal nutrition, egg quality and breeding success of Scottish Ptarmigan Lagopus mutus

Samples of Ptarmigan eggs were hatched in captivity. They had been taken from ground adjacent to study areas where the wild birds' breeding success by early August, and their food plants, were measured.
Almost all deaths of chicks in captivity occurred within a few days of hatching and examination postmortem showed no specific cause. The proportion of chicks which died in captivity before 15 days of age varied markedly between years and study areas. These variations occurred in parallel with variations in breeding success in the wild populations from which the eggs had been taken. It was concluded that the survival of chicks both in captivity and in the wild was determined in part before the eggs hatched.
Variations in breeding success from year to year were correlated with the number of days that the food plants had been growing before the hens finished laying. Breeding success in the wild and chick survival in captivity were better for a 'rich' area overlying some base-rich rocks than for a 'poor' one overlying granite and with less blaeberry. It was inferred that, as in Red Grouse, maternal nutrition affected breeding success through the quality of the eggs.     

Climate fluctuation effects on breeding of blue penguins (Eudyptula minor)

Abstract

El Niño and La Niña climate perturbations alter sea currents and food availability for seabirds in many areas of the world. This changes their breeding success and mortality. Blue penguin (Eudyptula minor) breeding success is dependent upon whether one or two clutches per season are laid, and the hatching and fledging success of these clutches. This study uses six years of data from five blue penguin breeding colonies, three from Taiaroa Head, Otago Peninsula and two from Oamaru, to examine whether annual variation in breeding success correlates with El Niño/La Niña perturbations. When La Niña conditions prevailed, penguins started breeding later, and there was a lower proportion of double breeders than in El Niño and normal years. The probability of a newly hatched chick surviving to fledging was also dependent on whether large‐scale climatic conditions prevailed, whereas hatching success and overall breeding success (number of fledged chicks per breeding pair) showed no correlation with climate perturbations.     

Breeding biology of the Barn Owl Tyto alba in central Mali

Data were obtained on 178 clutches of African Barn Owls in central Mali from four breeding seasons during 1979–1983. Significantly more clutches were laid in 1979–1980 and significantly fewer in 1980– 1981 than the average for the 4 years and there were significantly more clutches laid in the middle period of the annual breeding season. The egg volume was significantly smaller at the beginning of the breeding season and significantly larger in the middle than the overall mean with eggs of second clutches being larger than those of first clutches. The clutch size was 605 eggs of which 479 hatched. The number of young fledged per successful nest was 319 and was 1 83 for all nesting attempts. The month was the only variable shown to affect significantly the clutch size, eggs hatched and fledging rate, the highest success rates being associated with the middle of the breeding period. The average interval between the hatching of eggs was 2–31 days. Survival rates (47'1%) to fledging were significantly affected by year (1981–1982 being the least) and month (mid-season birds the best). The order of hatching significantly affected age at death or disappearance, the first-hatched birds surviving the longest. The year significantly affected age at fledging, the young from the year in which most clutches were laid leaving the nest at the youngest age and those associated with the year having the least number of clutches remaining in the nest the longest. The month of hatching also affected fledging age, birds at the extremes of the breeding season fledging at older ages. The discussion compares these data with those from elsewhere.