Philopatry and colony fidelity of Shags Phalacrocorax aristotelis on the east coast of Britain

Shag Phalacrocorax aristotelis colonies from the Firth of Forth south to the Fame Islands (SE Scotland and NE England) were surveyed in 1982 and 1983 for nesting Shags ringed as chicks or adults in the previous 20 years. On average, 5% of 863 Shags ringed as chicks were breeding away from their natal colony, and only 1% of 401 marked adults had left their breeding colony. The proportion p of chicks that settled at or beyond distance D from the natal colony followed the relationship p = 0.263D-^0.771, implying that more Shags moved 10 km or beyond to breed than expected from diffusion models or a constant-rate dispersal model; adjusted for birds that settled outside the study area, the relationship was p = 0.314D^-0771. There was no effect of colony size on emigration rate, but females showed a greater tendency to settle away from the natal colony than did males. At the natal colony, more males (69%) than females (47%) nested within 300 m of their birthplace. Two mechanisms appear to determine where a Shag settles to breed: a navigational one governing the return to the natal colony, plus the competitive process of nest-site establishment.     

Nest spacing in relation to settlement time in colonial cliff swallows

How colonial animals space their nests in relation to conspecifics may provide clues as to whether coloniality provides net benefits or occurs only because breeding sites are limited. We examined how nearest-neighbour distance varied in relation to settlement time in the highly colonial cliff swallow, Petrochelidon pyrrhonota, comparing observed nearest-neighbour distances to those expected if birds spread out to maximize nest spacing. Cliff swallows generally settled closer to each other than required by the available substrate, and clustered their nests closer in large colonies than in small ones. The first settlers at a colony site spaced themselves further apart than later arrivals but did not maximize nearest-neighbour distances. The first arrivals maintained greater nest spacing throughout the season than did birds that arrived later. Colony size and amount of nesting substrate had no effect on initial settlement distances of the first arrivals, but eventual nearest-neighbour distances declined with colony size. First arrivals may gain less from nesting with conspecifics and thus are less likely to cluster their nests than later arrivals, which may often be young or na?ve birds that gain more from the social benefits of colonial nesting. The results are consistent with the presumed social advantages cliff swallows receive from coloniality and do not support the hypothesis that colonies result from nesting site limitation. Copyright 2000 The Association for the Study of Animal Behaviour.     

Growth and decline of a penguin colony and the influence on nesting density and reproductive success

Colonial breeding is characteristic of seabirds but nesting at high density has both advantages and disadvantages and may reduce survival and fecundity. African penguins (Spheniscus demersus) initiated breeding at Robben Island, South Africa in 1983. The breeding population on the island increased in the late 1990s and early 2000s before decreasing rapidly until 2010. Before the number breeding peaked, local nest density in the areas where the colony was initiated plateaued, suggesting that preferred nests sites were mostly occupied, and the area used by breeding birds expanded. However, it did not contract again as the population decreased, so that nesting density varied substantially. Breeding success was related positively to the prey available to the breeding birds and negatively to local nest density, particularly during the chick-rearing period, suggesting a density-dependence operating through social interactions in the colony, possibly exacerbated by poor prey availability when the breeding population was large. Although nest density at Robben Island was not high, nesting burrows, which probably reduce the incidence of aggressive encounters in the colony, are scarce and our results suggest that habitat alteration has modified the strength of density-dependent relationships for African penguins. Gaining a better understanding of how density dependence affects fecundity and population growth rates in colonial breeders is important for informing conservation management of the African penguin and other threatened taxa.     

Breeding dispersal in a heterogeneous landscape: the influence of habitat and nesting success in greater snow geese

Despite numerous studies on breeding dispersal, it is still unclear how habitat heterogeneity and previous nesting success interact to determine nest-site fidelity at various spatial scales. In this context, we investigated factors affecting breeding dispersal in greater snow geese (Anser caerulescens atlanticus), an Arctic breeding species nesting in two contrasting habitats (wetlands and mesic tundra) with variable pattern of snowmelt at the time of settlement in spring. From 1994 to 2005, we monitored the nesting success and breeding dispersal of individually marked females. We found that snow geese showed a moderate amount of nest-site fidelity and considerable individual variability in dispersal distance over consecutive nesting attempts. This variability can be partly accounted for by the annual timing of snowmelt. Despite this environmental constraint, habitat differences at the colony level consistently affected nesting success and settlement patterns. Females nesting in wetlands had higher nesting success than those nesting in mesic tundra. Moreover, geese responded adaptively to spatial heterogeneity by showing fidelity to their nesting habitat, independently of snowmelt pattern. From year to year, geese were more likely to move from mesic to high-quality wetland habitat, regardless of previous nesting success and without cost on their subsequent nesting performance. The unpredictability of snowmelt and the low cost of changing site apparently favour breeding-site dispersal although habitat quality promotes fidelity at the scale of habitat patches.     

Movements of Eiders Somateria mollissima on the east coast of Britain

Movement patterns, sex differences in natal dispersal and breeding dispersal, and interchange of birds between colonies were studied in the population of Eiders Somateria mollissima breeding on the east coast of Britain. First-winter Eiders reared at the Sands of Forvie, Grampian, remain at or close to the colony, while most adults move about 100 km south to winter on the Firths of Forth and Tay. A proportion of the Forvie population is sedentary. Eiders which breed in Northumberland either move north to winter on the Tay and Forth estuaries or remain close to their breeding areas. Eiders breeding in Fife are sedentary. Recoveries of British-ringed Eiders in Scandinavia indicate that some British-born males join the Baltic breeding population, probably by pairing with Scandinavian females wintering in Britain. There is extensive natal dispersal of males from Forvie, with more than twice as many Forvie-bred females as males returning to the colony to breed. The breeding dispersal of males is also twice that of females. Dispersal of males from the relatively sedentary Forvie wintering population is less than that from the breeding population. Previous work suggested that at Forvie sedentary birds nesting close to the estuary were genetically isolated from migratory ones nesting along the coast. This situation is less clear cut than had been supposed previously, with many migrants nesting close to the estuary. It is unlikely that the genetic differences between females nesting in different parts of the Forvie colony will remain stable in the long term, due to the natal and breeding dispersal of males.     

FACTORS AFFECTING THE AGE OF FIRST BREEDING OF THE KITTIWAKE RISSA TRIDACTYLA

At a Kittiwake colony in Northumberland, 80% of those birds which returned to their natal colony to breed were males and these supplied 52% of all male recruits. More females breed away from their natal colony than males. There was no differences in the proportions of young fledged from sites in the centre or at the edge of the colony, or by parents of different experience, which returned to breed. Kittiwakes breed for the first time at ages from 3 to 8 years, but most at 4 or S years old. Males arrive back at the colony at an earlier age than females and breed for the first time one year earlier. Males obtaining sites at the centre of the colony first breed at an earlier age than those at the edges. Neither the age nor the area of first breeding appear to be transmitted from parent to offspring. Males breeding first aged 4 years or younger produced more young than those which first bred aged 5 years or older, despite their partners laying smaller clutches. This difference was most marked among those males recruited to sites in the centre of the colony. The advantage of this earlier breeding is counteracted by a lower survival rate among those males which start to breed at the younger ages. In all breeding Kittiwakes, annual reproductive output increases with experience while annual survival rates decrease. Once they had started to breed, many birds failed to breed in one subsequent season. Nearly 60% of these cases of intermittent breeding occurred in the year following first breeding. Intermittent breeding was most frequent among young birds and among females. It is suggested that each breeding involves a cost to the individual in terms of reduced survival, and that deferred and intermittent breeding are means of guarding survival. A model is proposed whereby the age at which a bird starts to breed, the nesting site which it obtains, and its subsequent breeding strategy result in each individual producing an optimal number of reproducing offspring in its lifetime, relative to its quality.     

Individual breeding decisions and long-term reproductive strategy in the King Penguin Aptenodytes patagonicus

Although studied for 35 years, knowledge of the reproductive biology of the King Penguin Aptenodytes patagonicus remains incomplete. The chick requires more than 12 months of care, which extends the breeding cycle, including moult, to more than one year, i.e. the King Penguin is neither annual nor biennial. In an attempt to resolve ambiguities in the literature and to elucidate the long-term breeding strategy of the species, we studied breeciing frequency at the individual level, considering the decision to breed in relation to breeding history over the previous few years. Although adult birds attempted to breed annually (0.83 breeding attempts per year), successful rearing occurred, at best, every two years only (maximum of 0.41 fledged chick per pair). Comparing successive years, the number of breeding birds in the colony was stable but the number of fledged chicks varied from 29 to 278 over eight years. These results suggest that King Penguins adopt (as individuals) an opportunistic reproductive strategy, in that they usually lay an egg every year, even when failure is certain. Nevertheless, the decision to breed was not entirely blind, and we identified groups of birds that invested differentially in breeding attempts. The decision to breed was related to the previous breeding frequency, i.e. 81% of the birds that had bred continuously in the past started a new breeding attempt, but only 67% of birds that had missed a year did so. In intermittent breeders, birds that had bred frequently, more often started a new breeding attempt than birds that had largely missed years (71% versus 57%, respectively). Classes of breeders could correspond to age classes, to birds of different breeding quality or to alternative breeding strategies coexisting in the species. Testing the hypothesis that reproductive effort increases with age should be possible in future.     

Assessing aural and visual cueing as tools for seabird management

Social attraction, that is, mimicking of active and productive colonies via audio playback of calls of breeding conspecifics and the use of decoys, is commonly used to attract birds to newly established or restored breeding sites. However, little is known about the relative importance of aural versus visual cues for identify nesting areas. Such information is important for design and evaluation of management protocols. We studied the effectiveness of decoys (visual cues) and playbacks (audio cues) as methods for restoring a colony of common terns (Sterna hirundo) at Muskeget Island, Massachusetts, USA. We used a 2-year, crossover experiment with 3 treatment areas: audio and visual, audio only, and visual only. We reversed treatment areas in the second year to control for previous nesting area or substrate preference. In both years, nests were built 9–101 m downwind of loudspeakers. There was no overlap in areas used for nesting between years and no nests were built within decoy plots in either year. Behavioral observations showed that birds responded to decoys only when within range of sound treatments. Conspecific vocalizations appear to be important proximate cues for seabird colony site selection and should be given priority in management protocols using social attraction. © 2011 The Wildlife Society.     

Estimating dispersal distributions at multiple scales: within‐colony and among‐colony dispersal rates,distances and directions in European Shags Phalacrocorax aristotelis

Knowledge of the rate, distance and direction of dispersal within and among breeding areas is required to understand and predict demographic and genetic connectivity and resulting population and evolutionary dynamics. However dispersal rates, and the full distributions of dispersal distances and directions, are rarely comprehensively estimated across all spatial scales relevant to wild populations. We used re‐sightings of European Shags Phalacrocorax aristotelis colour‐ringed as chicks on the Isle of May (IoM), UK, to quantify rates, distances and directions of dispersal from natal to subsequent breeding sites both within IoM (within‐colony dispersal) and across 27 other breeding colonies covering 1045 km of coastline (among‐colony dispersal). Additionally, we used non‐breeding season surveys covering 895 km of coastline to estimate breeding season detection probability and hence potential bias in estimated dispersal parameters. Within IoM, 99.6% of individuals dispersed between their natal and observed breeding nest‐site. The distribution of within‐colony dispersal distances was right‐skewed; mean distance was shorter than expected given random settlement within IoM, yet some individuals dispersed long distances within the colony. The distribution of within‐colony dispersal directions was non‐uniform but did not differ from expectation given the spatial arrangement of nest‐sites. However, 10% of all 460 colour‐ringed adults that were located breeding had dispersed to a different colony. The maximum observed dispersal distance (170 km) was much smaller than the maximum distance surveyed (690 km). The distribution of among‐colony dispersal distances was again right‐skewed. Among‐colony dispersal was directional, and differed from random expectation and from the distribution of within‐colony dispersal directions. Non‐breeding season surveys suggested that the probability of detecting a colour‐ringed adult at its breeding location was high in the northeastern UK (98%). Estimated dispersal rates and distributions were therefore robust to incomplete detection. Overall, these data demonstrate skewed and directionally divergent dispersal distributions across small (within‐colony) and large (among‐colony) scales, indicating that dispersal could create genetic and demographic connectivity within the study area.     

Breeding site fidelity and natal philopatry in the Redshank Tringa totanus

This paper presents the results of a study carried out on breeding Redshank in the Ribble Marshes, Lancashire, England.
Redshank tend to return to the same breeding area year after year. There was no detectable sex bias in return rates. Experienced birds were more site faithful than inexperienced birds, with previously successful birds exhibiting the highest degree of breeding site fidelity. Older, more experienced birds were more successful at hatching eggs than inexperienced birds.
Breeding dispersal was the same both within and between years. Faithful pairs and males nesting with a new mate dispersed significantly shorter distances than females nesting with a new partner. Dispersal distances in female Redshank were affected by breeding success: unsuccessful females, nesting with a new mate, dispersed significantly farther than successful females. A pair's breeding success influenced the following year's mate fidelity. However, other factors such as overwintering survival and date of return may also have influenced mate fidelity.
Redshank were highly faithful to their natal area; a high proportion of birds that survived post-fledging mortality returned to breed in their area of birth. No sex bias in natal dispersal was detected. Approximately 50% of Redshank breed in their first year of life.     

THE EFFECT OF AGE ON THE BREEDING BIOLOGY OF THE KITTTWAKE RISSA TRIDACTYLA

1. From 1954 to 1956 inclusive, the biology of individual marked Kittiwakes was studied at North Shields, Northumberland.
2. It was concluded that older Kittiwakes reacted to the breeding stimulus earlier, more intensively and with greater success than younger breeding birds.
3. Birds with previous breeding experience returned to the colony before birds breeding for the first time and these before non-breeders.
4. Before breeding started, birds which had bred previously spent more of their time at the colony than those about to breed for the first time.
5. Birds breeding for at least the second time laid the first egg 7·5 days earlier than those breeding for the first time.
6. Breeding started one day later for every four days the return to the colony was postponed.
7. Older breeding birds showed greater nest-site tenacity, laid larger clutches and had greater breeding success than younger birds.
8. The chicks in broods of two (but not of one) increased in weight more rapidly where the parents had previous experience.
9. Breeding Kittiwakes showed strong colony tenacity, but 24% of the marked non-breeding birds were subsequently seen in other colonies.
10. Over half the birds retained the same mate as in the previous year.     

Habitat selection responses of parents to offspring predation risk: an experimental test

The ability of nest predation to influence habitat settlement decisions in birds is widely debated, despite its importance in limiting fitness. Here, we experimentally manipulated nest predation risk across a landscape and asked the question, do migratory birds assess and respond to variation in nest predation risk when choosing breeding habitats? We examined habitat preference by quantifying the density and settlement date of eight species of migratory passerines breeding in areas with and without intact nest predator communities. We found consistently more individuals nesting in areas with reduced nest predation than in areas with intact predator assemblages, although predation risk had no influence on settlement or breeding phenology. Additionally, those individuals occupying safer nesting habitats exhibited increased singing activity. These findings support a causal relationship between habitat choice and nest predation risk and suggest the importance of nest predation risk in shaping avian community structure and breeding activity.     

American Kestrel reproduction: evidence for the selection hypothesis and the role of dispersal

Many hypotheses attempt to explain why younger, less experienced birds have a relatively low reproductive output. We evaluated reproductive patterns of marked American Kestrels Falco sparverius nesting in boxes in southwestern Idaho from 1992 to 2006 to test predictions of these hypotheses. Results were consistent with the selection (differential mortality) hypothesis and did not support the constraint, restraint or recruitment hypotheses. Most known-age Kestrels nested in their first year of life, and there was no apparent short-term or long-term reproductive advantage to delayed breeding. The number of years that Kestrels nested in study area boxes ranged from 1 to 6 years, with most Kestrels nesting in only 1 year. Reproductive rates were higher for birds with at least 1 year of nesting experience than for birds nesting in boxes for the first time. After 2 years of nesting, reproductive rates levelled off; there was no evidence for additional improvement or for senescence. Differences in reproductive output with experience/age were due to variation among and not within individuals. Individuals that nested in more than 1 year had similar reproductive rates in their first and second years. Poor producers either died or dispersed after 1 year of nesting in study area boxes. Successful females that nested early in the season and successful males that had been produced locally had the highest probability of returning to nest in a subsequent year.     

Barn swallows prefer to nest at sites hidden from neighboring nests within a loose colony

Some loosely colonial species scatter their nests within a colony. Most studies on mechanisms that generate such scattered nests within a colony have primarily focused on the measurement of distance related to the intensity of interactions among conspecific neighbors. However, these interactions can change not only with distance between nests but also with other factors. In particular, whether the nest is hidden from neighbors can strongly affect the intensity of interaction for birds that usually use sight for communication. To show the possible benefit of nesting at sites hidden from neighboring nests, we studied the nest arrangements and settlement patterns within a barn swallow colony. We tested whether the swallows prefer nesting at sites hidden from neighboring nests in cattle barns that are often used as colony sites by the swallows in Japan. We found that the sides of ceiling beams hidden from neighbors were preferred as nesting sites. A randomization test suggested that the rectangular sections occupied by swallows were not spaced out within a colony. The results suggested that the swallows preferred nesting sites hidden from their neighbors and did not space out their nests within a colony. This paper also refutes intraspecific brood parasitism as a plausible function of this behavior.     

THE BREEDING OF THE LESSER FLAMINGO IN THE MWERU WANTIPA, NORTHERN RHODESIA.

A hitherto unreported breeding site of the Lesser Flamingo Phoeniconaias minor is recorded in the Mweru Wantipa or Mweru Marsh, Northern Rhodesia. The site of the nesting colony, its structure, nest construction and behaviour of the birds is described. The importance of flooding in determining the site of the nesting colony is noted; adverse conditions caused by a rise in the water-level of Lake Mweru Wantipa appear to have completely destroyed the nests. The instability of such environments does not offer much chance of regular and permanent breeding, and may influence local movements on the part of the birds.     

Breeding dispersal and site-fidelity in three monogamous wader species in the Western Isles, U.K.

Adult and post-natal breeding-dispersal movements of Redshank Tringa totanus , Dunlin Calidris alpina and Ringed Plover Charadrius hiaticula were quantified in west Scotland. Data from 373 returning birds marked as breeding adults and 142 returning birds marked as chicks are presented. Unlike most previous studies, this study measured dispersal movements up to 40 km and attempted to overcome distance-related biases. For each species, adult males were significantly more nest site-faithful between years than were adult females. Likewise, first-time breeding males bred significantly closer to their natal site than did females. The settlement intensity per unit area of habitat showed marked differences between species, age classes and sex in the relative attractiveness of potential areas at different distances to settling birds. Adult dispersal was negatively and significantly related to breeding success in the previous year and positively related to capture on the nest in the previous year. Dispersal movements of adult Ringed Plover breeding on unstable cultivated habitats were significantly related to habitat quality (measured in terms of nest survival) and were always to habitat of better or equal quality.     

5. LIVINGSTONE IN NORTH-EASTERN RHODESIA

Sugg, M. St. J. 1974. Mensural and moult data from a breeding colony of Pied Kingfishers. Ostrich 45: 227–234.

Pied Kingfishers were ringed over two breeding seasons at a breeding colony on the Kenyan shore of Lake Victoria. Data were collected on weight, wing and bill length, injury and moult. Bill abrasion and breakage from nest excavation was found in both sexes and regeneration of worn and broken bills was recorded.

Adult birds of both sexes returned to the colony to breed but no juvenile was recaptured the year after hatching. Females are slightly larger than males (wing length) and both sexes showed a weight increase in the evening prior to roosting. Juveniles had shorter wings and bills than adults but their weights were similar. Their bills were short, soft, weak and the gape was salmon pink. No juveniles showed any moult. Adult moult records support the sequence and duration of moult suggested by Douthwaite (1971) though no overall decrease of moult activity was recorded during the breeding season.     

Sex, age, experience and condition as factors affecting arrival date in prospecting common terns, Sterna hirundo

Several studies have shown that seabird colonies consist to a large extent of young nonbreeders (prospectors). These individuals appear at the colony later in the season than established breeders. The reasons for this late arrival have remained unclear in most cases, mainly because of technical difficulties in collecting sufficient data from nonbreeding individuals. We used a novel transponder system to identify remotely the members of a common tern colony, including nonbreeders, during eight breeding seasons and we combined the system with automatic balances. Ninety-two per cent of prospectors returned for the first time when 2 years old and 88.9% of recruits to the breeding population had spent at least one previous season at the colony as prospectors. In both sexes, most individuals prospected for one season, but more males than females prospected for more than one season, although a higher proportion of females started breeding without a previous prospecting phase. Terns arrived earlier in the season the older they were and the more experience of the colony they had, but experience proved to be more important than age. Prospectors gained about 3 weeks with a previous prospecting season whereas an additional year of age allowed birds to arrive only about 6 days earlier. Prospectors returning later in the season arrived with lower body masses. Males on average arrived earlier at the colony than females. Copyright 2003 Published by Elsevier Science Ltd on behalf of The Association for the Study of Animal Behaviour.      

Effects of vegetation on nest microclimate and breeding performance of lesser black-backed gulls (Larus fuscus)

For birds that breed in large colonies, the overall area occupied by the colony generally comprises several sub-areas that differ in physical and social features such as vegetation and breeding density. Birds arriving at a breeding colony select their nesting sites through a hierarchical process of selecting a sub-area, then a particular nest site with appropriate biotic and physical attributes. Optimal vegetation cover is one such important attribute. Many ground nesting gulls preferentially select nest sites that provide shelter during reproduction, but this presumably has to be balanced against any costs such as reduced visibility of potential predators. The effects of vegetation height in the sub-areas within a colony, and of the amount of vegetation in the immediate vicinity of the nest on nest microclimate were investigated in lesser black-backed gulls Larus fuscus in a colony in which overall vegetation height differed in different sub-areas and was patchily distributed within these areas. Tall vegetation did have a sheltering effect, and this was positively related with chick growth. However, this vegetation area was associated with lower breeding densities, relatively late laying birds and lower chick survival rate, suggesting that sub-areas with tall vegetation held more lower-quality or young breeders. Within the sub-areas, the birds preferentially selected nest sites with more surrounding vegetation, and this was positively correlated with their hatching success.     

The post-fledging survival of young Guillemots Uria aalge in relation to hatching date and growth

The post-fledging survival of a total of 1277 young Guillemots Uria aalge ringed in 6 years was assessed using sightings of 267 individuals back at the natal colony and recoveries of 46 ringed birds. In two years there was a significant decline in survival prospects with estimated hatching date, the first time such a trend has been demonstrated in the Alcidae. In these years pairs breeding early had a 2–3 times greater chance of having a young survive to return to the colony than pairs breeding three weeks later. Hatching date had no effect on survival prospects in the other four seasons and in none of the years did chick body condition have a demonstrable effect on post-fledging survival.