BREEDING PERFORMANCE OF PUFFINS FRATERCULA ARCTICA IN RELATION TO NEST DENSITY, LAYING DATE AND YEAR

The paper presents data on the breeding and predation of Puffins in two areas of different nest density within a single colony on Dun, St Kilda group, Outer Hebrides in 1973-78.
Within a season birds laying early had a slightly higher nesting success than birds laying late, but laying date had little influence on the peak and fledging weights of young. The main disadvantage in late laying was a reduced chance of relaying if the first egg was lost.
Breeding success and chick weights varied from year to year. The 1974 season was the least successful with the lowest nesting success, lowest frequency of feeds, lowest calorific value of feeds, lightest chicks and slowest growth. Overall breeding performance was not related to the annual mean laying dates.
In all years pairs nesting in the area of high nest density did better than pairs nesting at low density. The effect is attributed to differential predation and disturbance by predatory gulls. At least 4.2% of adult Puffins breeding in the area of low burrow density were killed by gulls each breeding season; this is higher than the total annual mortality found in three other studies. Only 0.9% of adults from the high density area were found killed. The subpopulation in the low density area cannot survive without much immigration, yet there is no evidence that this happens.     

THE BREEDING OF CORY'S SHEARWATER CALONECTRIS DIOMEDEA ON THE SALVAGE ISLANDS

The breeding of Cory's Shearwater on Selvagem Grande was studied during the seasons of 1968 and 1969, on a number of specially timed visits. After a pre-laying exodus (duration not determined) the birds returned in a mass to the island on 26 May 1969 and laying began immediately reaching its peak on 31 May. Egg dimensions and weights are tabulated. In most cases the female handed over to the male immediately after laying, and the two sexes incubated in alternating spells averaging about six days. The incubation period averaged 53-8 days. The chicks reached their maximum weight at about 53 days, but the fledging period was not determined. In 1968 young birds were leaving the nest on 22–25 October, about 90 days after the 1969 mean hatching date. Herring Gulls were important egg predators, but losses of chicks were few, 29 out of a sample of 30 being alive at the age of about 60 days.     

Survival of Herring Gull Larus argentatus chicks: an experimental analysis of the need for early breeding

In 1985 and 1990, we manipulated the hatching date of Herring Gulls Larus argentatus by exchanging complete clutches between pairs which had started egg laying on different dates in the season. The aim was to investigate whether the observed seasonal decline in Hedging success can be explained by laying date. In both years, fledging success of advanced pairs followed the natural seasonal trend, supporting the date hypothesis. However, in both years, Hedging success of delayed pairs remained as high as that of control pairs with the same original hatching date as the foster parents, supporting the parental quality hypothesis. These data suggest that the delayed pairs may have compensated for less favourable environmental conditions. Although conspecific predation was the main direct cause of chick mortality, food shortage probably indirectly determined the mortality rate. A model of fledging success in relation to the timing of egg laying predicts that when food is abundant the optimal hatching date coincides with the mean hatching date of the colony (synchronization advantage). When food is less abundant, early breeders have the highest fledging success.     

FACTORS AFFECTING BREEDING OF RAZORBILLS ALCA TORDA ON SKOKHOLM

A study of the breeding biology of the Razorbill was carried out on Skokholm (South Wales) during 1971-73. Birds ringed or colour ringed before the study began provided additional information upon the effects of age on breeding. Mean laying date was delayed in 1972, compared with 1971; the effect is attributed chiefly to stormy weather which upset colony attendance. Eggs were also smaller in 1972. A seasonal decline in egg size (volume) was noted in all three years, attributed mainly to the later laying of young birds. Egg size increased with age, at least up to the fifteenth year. Eggs lost totalled 30% of those laid; 73% of this total was due to predation by Herring Gulls and of Jackdaws. Most losses (45%) occurred during the first 10 days after laying. Of lost eggs, 25% were replaced, usually 14 days after the loss of the original; only eggs laid and lost early in the season could be replaced. Only 7% of the chicks which hatched failed to fledge. Most (62.5%) chick losses occurred in the first week of nestling life, when chick weight was related to egg size. Afterwards, both growth rate and fledging weight were independent of egg size. The chicks fledging early in the season were heavier than later chicks. Failure to fledge was mainly due to a breakdown in behaviour between parent and young, rather than to predation. Breeding success was highest for birds breeding early in the season, most of which were older, more experienced breeders. These laid early enough to replace an egg if it was lost; they produced large eggs, and their chicks were therefore both heavier than average during the critical first 7–10 days of life, and fledged at a high weight. Thus experience accumulated with age, and the ability to lay early in the season are important for successful breeding in the Razorbill.     

Habitat selection and breeding success in Yellow-legged Gulls Larus cachinnans

The habitat selection and breeding performance of Yellow-legged Gulls Larus cachinnans were studied in the Medes Islands colony, northeastern Spain, during 1995 and 1996. Of the three main habitats on the islands (shrubs, grass and bare areas), gulls first occupied those with the highest percentage of tall vegetation. Gulls tended to select nest sites with 20–75% cover despite great differences in the cover in the habitats and territories, suggesting that the presence of a suitable nest site may play a major role in the choice of breeding habitat. Nest-site tenacity did not influence the preferences of gulls at any level since the same pattern of choice was observed in an area subjected to annual culls (i.e. where most of the breeding pairs were culled annually and replaced by naive birds). In spite of great differences in the physical characteristics of the habitats, little difference was found in breeding performance of the gulls between habitats. Gulls nesting in the least preferred habitat (i.e. mainly bare) had smaller clutches than those nesting in the other two habitats, possibly as a result of their later seasonal laying. Despite the similar breeding success in different habitats, gulls did not seem to distribute according to the ideal free model reported for Herring Gulls Larus argentatus since the density in the preferred habitat (i.e. shrubs) was never higher than in the other two. We suggest that the habitat selection by Yellow-legged Gulls within the colony could follow an ideal despotic distribution.     

A comparison of Herring Gull Larus argentatus and Lesser Black-backed Gull Larus fuscus nest sites: their characteristics and relationships with breeding success

Nest sites used by Herring Gulls and Lesser Black-backed Gulls in a mixed colony were compared. Correlations between features surrounding a nest and breeding success at different stages of the breeding cycle are presented. Lesser Black-backed Gulls nested on more vegetated and flatter areas than did Herring Gulls, even though the latter species had a lower hatching success at less vegetated sites. The difference in the general topography of nest sites between the species suggests that the Lesser Black-backed Gull, through an individual defence strategy, may be better adapted to use sites which are accessible to ground predators than the Herring Gull. The fledging success of Lesser Black-backed Gulls tended to increase with an increased proportion of relatively tall vegetation close to the nest. The Lesser Black-backed Gull may therefore be able to leave its young unattended and rely on their concealment for protection against predators. In contrast, the Herring Gull may rely more on parental vigilance to protect young. More frequent attendance by adult Herring Gulls at the nest site during chick rearing compared with Lesser Black-backed Gulls supports this hypothesis.     

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.     

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.     

TIMING AND SPACING OF BROODS IN THE BLACK-HEADED GULL LARUS RIDIBUNDUS

The nests of the Black-headed Gull Larus ridibundus are closely aggregated into dense colonies and their use synchronized, these two phenomena together tending to produce a maximal clumping effect. Within such a colony however, nests were found to be spaced out to produce a non-random uniform distribution. The commonest distance between neighbouring nests was found to be about one metre, in contrast to related species. This study was concerned with two aspects of this distribution pattern; its survival value and its behavioural causation.
It was found that pairs nesting just outside the colony had a much lower breeding success than those nesting in the colony and that nests on the colony fringe had a slightly lower success than those in the centre. Pairs laying during the peak laying period had a higher breeding success than pairs laying either earlier or later in the season. Since by far the most important mortality agent was predation, it seems likely that both clustering and synchronization of nesting function as antipredator systems and arguments in favour of this are discussed.
Variations in nest-spacing within the colony were not correlated with variations in breeding success.
In the causation of the spacing between nests, territorial aggression was demonstrated to be an effective dispersion mechanism and the way in which this mechanism works was investigated in detail.
This spacing mechanism was not sufficient by itself to explain the observed densities, which were higher than one would expect from the aggression alone; there was also some tendency for birds establishing a new nest-site to cluster close to others. The interaction between this, the territorial aggression of the residents and the subsequent avoidance responses of the settling birds, can explain the nest spacing pattern and probably also the observed densities.     

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.     

Effects of the parasitic botfly Philornis carinatus on nestling house wrens,Troglodytes aedon,in Costa Rica

I studied the life cycle of a botfly (Diptera: Muscidae: Philornis carinatus) and examined the effects of botfly ectoparasitism on nestling house wrens (Passeriformes: Troglodytidae: Troglodytes aedon) during three years in Costa Rica. At three study sites, I found that nestlings were relatively unaffected by botflies, in contrast to all other studies of birds infected with philornid botflies. At Monteverde, the main study site, infected chicks grew slightly slower and had slightly shorter tarsi and wing chords than uninfected chicks, but both groups fledged at similar weights. Since weight at fledging is the only growth character associated with post-fledging survivorship, botfly infections likely cost wrens little in terms of fitness. At all sites, fledging success did not differ between infected and uninfected nests. Botfly infections were more prevalent at two lower elevation sites than at the high elevation Monteverde side. Infection prevalence increased during the nesting season at all study sites, which suggests a botfly life cycle in which adult population levels increase during the wren breeding season and then decline during a dormant period when wrens are not nesting. Finally, botflies may attack chicks throughout the period before fledging, but there is no indication they locate nests before hatching. In sum, botfly parasitism on wrens appears to be benign, perhaps because the study sites are at the edge of the botfly's range or because wrens are not a preferred host.     

PARENTAL QUALITY AND SELECTION ON EGG SIZE IN THE MAGELLANIC PENGUIN

We examined the relative contributions of egg size and parental quality to hatching success, fledging success, and chick growth in the Magellanic penguin (Spheniscus magellanicus) be exchanging clutches between nests to reduce the covariation between egg and parental factors. Among control nests, fledging success increased slightly with egg size. However, the effect of egg size independently of parental quality was limited to an influence on chick mass and size for the first 10 days post-hatching. In contrast, attributes of the parents influenced nesting success and chick size at fledging, independently of the egg size actually raised. We suggest that the common occurrence of a positive phenotypic correlation between egg size and fledging success is due to two factors: (1) adults laying large eggs tend to be of higher quality; and (2) to the extent that egg size does influence early survival independently of parental quality, the effect on survival is due to a maternal effect on egg composition rather than an inherent effect of egg size.     

NESTING OF THE VILLAGE WEAVER PLOCEUS CUCULLATUS

From August to October 1975, 31 nesting colonies of Village Weavers were identified within an area of 3000 km², south of Lake Chad in Africa. The number of nests and the productivity of each colony were estimated by means of several samplings made during the course of the nesting season. Differences between bush colonies and village colonies are not relevant. The average length of the nesting season is 70 days. In each colony the number of nests increases during the first month and then decreases. The average number of fledging attempts is four per colony, with a period of about ten days from one to another. On a regional scale, breeding periods and fledging attempts are well synchronized. Each nest carries an average of 2.4 eggs and 1.9 young, the difference occurring only at hatching. Due to regional synchronism, the period of most intensive reproductive activity, at which the number of nests reaches its maximum, can be determined.     

Sex,growth rate,rank order after brood reduction,and hatching date affect first‐year survival of long‐lived Herring Gulls

Among most species of birds, survival from hatching throughout the first year of life is generally lower than subsequent survival rates. Survival of young birds during their first year may depend on a combination of selection, learning, unpredictable resources, and environmental events (i.e., post‐fledging factors). However, knowledge about post‐fledging development in long‐lived species is usually limited due to a lengthy immature stage when individuals are generally unobservable. Therefore, pre‐fledging characteristics are often used to predict the survival of young birds. We assessed effects of nestling growth rates, hatching date, hatching asynchrony, brood size and rank order after brood reduction, and sex on first‐year survival of 137 fledglings using a mark‐resighting analysis. We found that the survival probability (Φ_1yr = 0.39) of first‐year Herring Gulls (Larus argentatus) in our study colony located at the outer port of Zeebrugge (Belgium) was lower than that of older individuals (Φ_>1yr = 0.75). All 10 models best supported by our data included nestling growth rate, suggesting that variability in first‐year survival may be linked primarily to individual variation in growth. First‐year survival was negatively correlated with hatching date and rank order after brood reduction. Hence, carry‐over effects of breeding season events such as timing of breeding, early development, and social status had an influence on survival of Herring Gulls after fledging. Furthermore, we found sex‐biased mortality in first‐year Herring Gulls, with females (Φ_1yr = 0.45) surviving better than males (Φ_1yr = 0.38). Although adult survival is generally regarded as the key parameter driving population trajectories in long‐lived species, juvenile survival has recently been acknowledged as an important source of variability in population growth rates. Thus, increasing our knowledge of factors affecting age‐specific survival rates is necessary to improve our understanding of population dynamics and ultimately life‐history variation.     

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.     

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.     

The effect of body condition on the timing and success of breeding in Little Penguins Eudyptula minor

We studied the effect of parental body condition on the breeding biology of Little Penguins Eudyptula minor . Daily attendance patterns and body mass were recorded using an Automated Penguin Monitoring System, which collected arrival and departure masses for approximately 200 breeding birds over the 2000 and 2001 breeding seasons. Breeding success varied between the two years; 2000 was a year of average breeding success (fledging 1.07 chicks per pair), and 2001 a year of poor breeding success (fledging 0.53 chicks per pair). In both years, adult body condition (body mass divided by flipper length) increased significantly prior to laying. The laying period began over a month later in 2000 than in 2001, and birds in 2000 exhibited significantly better body condition at laying. However, the mean laying dates in 2000 were less variable than in the 2001 breeding season. Body condition appeared to influence both the time of breeding and breeding synchrony in Little Penguins. Breeding success was correlated with adult body condition at incubation in 2000 but not in 2001, indicating that success was not solely influenced by adult body condition at incubation.     

SELECTION OF COLONY SITES AND NEST SITES BY COMMON TERNS STERNA HIRUNDO IN OCEAN COUNTY, NEW JERSEY

Common Terns nested on 34 of 259 saltmarsh islands along 46 miles of Ocean County coastline, New Jersey. They nested on low islands of Spartina with less than 12%, windrow ranging in size from 0.6 to 108 acres. All islands selected by terns faced at least two miles of open water from at least one direction. Selection of islands seemed a response to vegetation, size, distance to the nearest island, distance to the nearest shore and exposure to open water. Of the 225 islands without nesting terns, only three fulfilled appropriate criteria. Eighty per cent of nests were situated on windrow. Terns laid larger clutches on windrow compated to those nesting in Spartina. Nearest neighbour distance on islands with low predation varied from 85 to 485 cm and was a function of space, vegetation, and the size of the island. Terns nested closer together on windrow compared with Spartina. Considering all colonies, the mean nearest neighbour distance negatively correlated with the number of nests. Skimmers, Oystercatchers, Laughing Gulls, and Herring Gulls nested in some of the tern colonies. All tern colonies on islands also occupied by nesting Herring Gulls suffered over 10% egg-loss by predation. Proportionally more nests were preyed upon in windrow than in Spartina. Clutch sizes in nests in windrow on islands with predation were significantly lower than those in windrow on islands without predation, even though clutch sizes in Spartina were similar on islands with and without predation. Flooding by exceptional high tides destroyed significantly more nests in Spartina than in windrow. The nests that survived in Spartina were built deeper, and their rim heights were higher than nests destroyed by tides. Choice of colony and nest sites is discussed in terms of the balance of two selection pressures: predation and tidal flooding. Tern nests on windrow are more exposed to predators, but safer from flooding, whereas those nests in Spartina are more susceptible to flooding and less vulnerable to predators. The recent invasion of Herring Gulls into Ocean County has significantly increased the rate of nest predation among affected colonies of Common Terns. It is possible that in future years the terns' behaviour may be modified in response to this new selective pressure.     

THE PROCESS OF COLONY FORMATION AMONG HERRING GULLS LARUS ARGENTATUS NESTING IN NEW JERSEY

We examined the pattern of colony occupation and egg-laying in five colonies of Herring Gulls nesting in New Jersey, U.S.A. Colonies formed from epicentres located in sparse bushes. The number of epicentres related to the number of birds nesting in the colonies. Colonies of over 250 pairs had more than one epicentre, whereas those with under 250 pairs had only one epicentre. Epicentres were not always in the geographical centre of the colonies. New territory-hunting pairs filled in the epicentre areas, and then nested outside these areas. The egg-laying pattern followed the settling pattern, but was more synchronous than the settling pattern. There was greater synchrony of egg-laying within sub-areas of the colonies than in the colonies as a whole. Further, synchrony correlated with the number of nests in sub-areas.     

Intraspecific nest parasitism in the grey starling (Sturnus cineraceus)

We studied intraspecific nest parasitism in the grey starling (Sturnus cineraceus) in 1992 and 1993. We used three criteria to detect nest parasitism: (i) the appearance of more than one egg per day while the host was laying; (ii) the appearance of extra eggs after the host completed its clutch; and (iii) the appearance of eggs which were of a different shape, size and color to other eggs in the clutch. There were 290 nests (157 nests in 1992; 133 nests in 1993) in which the clutch was completed early (clutches initiated before May 10). Twenty-nine (1992) and 32 (1993) nests contained at least one parasitic egg. Parasitic eggs hatched if they were laid during the laying period and early in the incubation period of their host, and a few of them fledged. Fledging success of parasitic eggs was not different from that of eggs in non-parasitized nests if parasitic eggs were laid during the host's laying period. However, fledging success of all parasitic eggs was fewer than that of eggs in non-parasitized nests. By comparison, fledging success of parasitized nests was not a great as that of non-parasitized nests.