Influence of food availability on demography and local population dynamics in a long-lived seabird

Few studies have addressed the effects of food availability as a proximate factor affecting local adult survival in long-lived organisms and their consequences at local population dynamics. We used capture-recapture analysis of resightings of 10 birth cohorts of ringed Audouin's gulls, Larus audouinii, to estimate adult survival and dispersal (both emigration and immigration). For the first time, permanent emigration (the transient effect in capture-recapture analysis) was modelled for the whole population and not only for the newly marked birds. Gulls exploit to a large extent fishes discarded from trawlers, and a trawling moratorium established since 1991 has decreased food supply for the colony. This was used as a natural experiment of food availability to assess its effects on adult survival and emigration. These and other demographic parameters were used in a projection modelling to assess the probabilities of extinction of the colony under two scenarios of lower and higher food availability. Food availability (together with the age of individuals) influenced emigration probabilities, but not adult survival, which was estimated at 0.91 (s.e. = 0.02). When food was in shorter supply during the chick-rearing period, emigration was very high (ca. 65%) for younger breeders, although this rate decreased sharply with age. Probabilities of extinction were very high when food availability was low, and when environmental stochasticity was introduced, and only stochastic immigration from the outside seemed to prevent extinction. The results highlight the importance of dispersal processes in the population dynamics of long-lived organisms.     

Survival in king penguins Aptenodytes patagonicus: temporal and sex-specific effects of environmental variability

We investigated annual adult survival rates of king penguins Aptenodytes patagonicus breeding at South Georgia during 6 years in relation to age/breeding experience, sex, and food availability. During the first 3 years of the study, when food availability was good, survival was 97.7% for experienced breeders, which confirmed the very high survival rates observed in penguins in general. In these years survival did not differ between the sexes, presumably because parental investment is shared equally between the sexes, and the sexual dimorphism is small in king penguins. Survival was lower for young, first-time breeders (83.0%). In experienced birds the annual survival rate decreased to 68-82% following a catastrophic year when food availability was extremely low. We address the question how parents balance their current investment in offspring against their chances to reproduce in the future. We argue that the high mortality rate among breeding individuals after the year of food stress provides support for previous suggestions that the response to increased costs in seabirds might be complex to predict and does not always follow intuitive expectations according to general life-history theory. We also found that females survived significantly less well than males following the bad year. We explain this result as follows: the male-biased sex ratio (56:44) that we observed in our study colony clearly does not result from lower female survival during normal conditions. An already existing skewed sex ratio forces males to delay the onset of breeding because of a lack of breeding partners. This in turn causes breeding females to be, on average, younger and less experienced than males and to have lower survival following a year of food shortage. In this study survival was linked with food availability and we suggest that this was connected to climatic/oceanographic features, such as the position of the Antarctic Polar Front Zone. We could, however, not verify this by anomalies in sea surface temperature data.     

Spatial contraction in a large gull colony in relation to the position of arctic fox dens

Many species of colonial ground-nesting birds are known to be sensitive to predation by terrestrial predators. The response of prey under the pressure of predation can either be direct (depletion of prey) or indirect due to prey avoiding the predator. We studied the recolonization of arctic foxes into a large and growing breeding colony of lesser black-backed gulls. The gull colony reached the size of 20,000 pairs during the period of no foxes. The number of breeding gulls continued to increase after the colonization of foxes and reached a maximum of 40,000 pairs about 15 years later. During the same time period, the spatial coverage of the gull colony shrunk from 31.4 down to 18.6 km², and the change in distribution was closely linked with position of fox dens, which ranged in number from one to three annually. In 2005, the number of breeding gulls decreased slightly with an ongoing shrinkage in spatial coverage. Food analyses from fox scats and stomachs showed that birds of the order Charadriiformes, particularly gulls, were the key prey item. Survival rates of gull nests were higher in the middle of the colony than at the colony edge close to a fox den. The colony area lost could be explained solely by the number of eggs removed by foxes but is unlikely, e.g., due to reclutching. We suggest that intraspecific predation contributes additionally to the effects of direct fox predation although to an unknown degree.     

Balancing Energy Budget in a Central-Place Forager: Which Habitat to Select in a Heterogeneous Environment?

Foraging animals are influenced by the distribution of food resources and predation risk that both vary in space and time. These constraints likely shape trade-offs involving time, energy, nutrition, and predator avoidance leading to a sequence of locations visited by individuals. According to the marginal-value theorem (MVT), a central-place forager must either increase load size or energy content when foraging farther from their central place. Although such a decision rule has the potential to shape movement and habitat selection patterns, few studies have addressed the mechanisms underlying habitat use at the landscape scale. Our objective was therefore to determine how Ring-billed gulls (Larus delawarensis) select their foraging habitats while nesting in a colony located in a heterogeneous landscape. Based on locations obtained by fine-scale GPS tracking, we used resource selection functions (RSFs) and residence time analyses to identify habitats selected by gulls for foraging during the incubation and brood rearing periods. We then combined this information to gull survey data, feeding rates, stomach contents, and calorimetric analyses to assess potential trade-offs. Throughout the breeding season, gulls selected landfills and transhipment sites that provided higher mean energy intake than agricultural lands or riparian habitats. They used landfills located farther from the colony where no deterrence program had been implemented but avoided those located closer where deterrence measures took place. On the other hand, gulls selected intensively cultured lands located relatively close to the colony during incubation. The number of gulls was then greater in fields covered by bare soil and peaked during soil preparation and seed sowing, which greatly increase food availability. Breeding Ring-billed gulls thus select habitats according to both their foraging profitability and distance from their nest while accounting for predation risk. This supports the predictions of the MVT for central-place foraging over large spatial scales.     

ECTOPARASITE LOAD IN A LAUGHING DOVE WITH A DEFORMED MANDIBLE

At Dyer Island, South Africa, observations of predation of seabirds by Cape Fur Seals Arctocephalus pusillus pusillus were made during 2004 and 2006/07. It was estimated that seals killed about 7% of adult African Penguins Spheniscus demersus annually. This may have contributed to the penguin colony not increasing in spite of food becoming more available in its vicinity during an eastward shift of epipelagic fish resources off South Africa at the start of the twenty-first century. Adult penguins were most susceptible to mortality in the breeding season as they returned to feed chicks in the evening. Seals also killed substantial numbers (3–9%) of Cape Cormorant Phalacrocorax capensis fledglings as they left the island, mostly in the morning. The losses of Cape Cormorant fledglings accounted for about 95% of the overall mortality of seabirds attributable to seals. Mortality inflicted by seals on adults of four species of cormorant was negligible and no predation by seals on gulls and terns was observed.     

Management-Induced Reproductive Failure and Breeding Dispersal in Double-Crested Cormorants on Lake Champlain

ABSTRACT We studied breeding dispersal of double-crested cormorants (Phalacrocorax auritus) associated with management practices that suppressed their reproduction on Lake Champlain in the northeastern United States. We implemented an experiment on one colony by spraying corn oil on cormorant eggs in portions of the colony and leaving other portions untreated. Gulls (Larus spp.) consumed cormorant eggs during the oiling process, but we reduced and then eliminated predation levels after the first year of the study. We used mark-recapture techniques within the experimental framework to measure rates of breeding dispersal for cormorants from the experimental colony and an unmanaged colony in Lake Champlain. Egg oiling increased the movement rate to the unmanaged colony by 3% during the year with no egg predation by gulls. When gulls depredated cormorant eggs at high rates during egg oiling, movement to the unmanaged colony increased by 20%. When cormorants are managed to reduce population sizes, methods that limit dispersal away from the managed colony may be most effective. Such methods would mitigate effects to nontarget populations and allow for a greater portion of the metapopulation to be managed.     

Determinants of local recruitment in a growing colony of Audouin's gull

1. Local recruitment of Audouin's gull ( Larus audouinii Payraudeau) was studied between 1988 and 1997 at the Ebro Delta colony (north-western Mediterranean). Since its establishment in 1981, the colony has dramatically grown to include, in 1997, 65% of the total world population. Several hypotheses were tested, involving the effects of a badger predatory event in 1994, and sex, age and cohort (year of birth) on recruitment.
2. Results supported the prediction that colony size influenced recruitment: the probability for any individual to have previously bred increased throughout the study, together with colony size. At the end of the study, 90% of breeders aged 4 years had already been recruited at age 3, the age of first reproduction by Audouin's gulls. As expected by the dramatic increase of breeding numbers, most local recruitment occurred at very young ages, especially when compared with other Laridae.
3. Neither food availability nor reproductive success affected recruitment. Recruitment was not affected by high nest predation by the badger, although after the event, the proportion of Ebro Delta birds nesting on the nearby Columbretes Islands tripled.
4. Probability of first reproduction depended on age: it was the highest at ages 3 and 4, and then decreased sharply with age to stabilize beyond age 6 to a value depending on the year and cohort but always very low (< 5%). Cohort and sex did not influence local recruitment.
5. Annual resighting rates ranged between 35% and 82%, and were higher for females. This may represent a sex-dependent suspension of breeding, probably as a trade-off between early recruitment and future survival.     

Breeding biology of Sabine’s gull (Xema sabini) in the Canadian high Arctic

The Sabine’s gull (Xema sabini) is a small seabird that breeds in select locations across the circumpolar Arctic, but there have been few studies on its breeding biology, particularly from the high Arctic. We studied nesting phenology, breeding effort, and breeding success of Sabine’s gulls over 5 years at a colony on a small island (Nasaruvaalik) in the Canadian high Arctic. Compared to studies in the low Arctic, nest initiation dates and adult body mass were more consistent across years, and reproductive success was typically higher at Nasaruvaalik Island. These differences may be related to the more predictable food sources available in the nearby polynya upon arrival from migration, as well as the lower predation pressure at our site.     

Avian life history variation along altitudinal gradients: an example with cardueline finches

Elevation has long been considered a major influence on the evolution of life-history traits. Most elevation-induced variation in life history traits has been attributed to changes in climate, duration of breeding season, predation, and food limitation. I use a phylogenetic approach to show that life histories are closely associated with breeding elevation in extant cardueline finches. Finches at high elevations had smaller clutches, fewer broods, and longer incubation periods. Neither food limitation nor nest predation appear to readily account for this strong elevational variation in cardueline life histories. However, juvenile survival may be greater at higher elevations as a result of prolonged parental care and shorter natal dispersal and can potentially compensate for reduced fecundity in high-elevation finches. Received: 28 September 1996 / Accepted: 24 March 1997     

Effect of interannual variations in sea-surface temperature on egg-laying parameters of black-tailed gulls (Larus crassirostris) at Teuri Island, Japan

Sea-surface temperature (SST) directly and indirectly affects the distribution and abundance of prey species for seabirds, so we expect variation in SST to be associated with variation in seabird life history traits. In black-tailed gulls (Larus crassirostris) at Teuri Island in northern Hokkaido, Japan, we investigated the diet of the gulls prior to egg laying in 2004 and 2005, and examined the influences of SST in March or April, when the gulls congregate in the colony, on egg-laying parameters using 13 years of data (1992–2004). The gulls fed on krill (Thysanoessa inermis) and fish prior to the egg laying. Mean first egg dates and clutch sizes were significantly and quadratically related to SST anomalies in March, but were not influenced by SST anomalies in April. There was no significant effect of SSTs in either March or April on egg volume. Sea-surface temperature anomalies in March of the years of early laying (−1 to 1°C) were higher than those in 2001 (−2.2°C), but lower than those in 1992 (+1.2°C) and 2004 (+1.1°C). Thysanoessa inermis congregates to spawn at the sea surface, when SSTs rise 3−4°C. Thus, a mismatch between food availability and the timing of egg production in the gulls could have occurred in these 3 years. This study suggests that SST fluctuations prior to laying are important in breeding success of black-tailed gulls.     

Interspecific kleptoparasitism in Audouin's Gull Lams audouinii at the Ebro Delta, northeast Spain: a behavioural response to low food availability

Kleptoparasitism of Audouin's Gull Lams audouinii on other seabirds, a foraging behaviour previously unrecorded for the species, was studied in the Ebro Delta, northeast Spain, during 1992. The relationship between kleptoparasitism and trawler activity was assessed through changes in the availability offish discards brought about by a trawling moratorium. When trawlers are not active, Audouin's Gulls use different feeding strategies, such as interspecific kleptoparasitism. Audoin's Gulls appeared to prefer to kleptoparasitize terns rather than gulls, but the success rate on gulls was higher. Most of the attacks rather than gulls, but the success rate on gulls was higher. Most of the attacks were performed by single Audouin's Gulls, although success per attack improved with group size. During the trawling moratorium, the existence of large seabird colonies in the Ebro Delta acted as a buffer for Audouin's Gull through kleptoparasitism, compensating partly for the reduced food supply.     

Breeding dispersal strategies following reproductive failure explain low apparent survival of immigrant Temminck's stints

In some animal populations, immigrants have lower survival than philopatric individuals. Costs of dispersal or low phenotypic quality of dispersers may explain the pattern. However, apparent adult survival estimates, which describe real survival combined with site fidelity cannot be separated from permanent emigration. Thus, heterogeneity in breeding dispersal propensities of immigrants and philopatrics can bias fitness correlates of dispersal. Differences in breeding dispersal propensities may be caused by different strategies in response to environmental cues inducing dispersal, such as reproductive success. In such cases, the reported differences between immigrants and philopatric individuals may not reflect true variation in survival. We studied whether dispersal status specific apparent adult survival is associated with reproductive success in a Temminck's stint Calidris temminckii population. We analysed two long term capture–recapture datasets characterised by low and high nest predation levels. Philopatric individuals had higher apparent adult survival than immigrants in both datasets and the difference was highlighted during the high nest predation period. By contrasting return rates between successful and unsuccessful breeders as a proxy for dispersal, we found that unsuccessful immigrants breeding for the first time dispersed more likely than successful immigrants, but such a pattern was not found among philopatric individuals. Our results support the hypothesis that immigrant and philopatric individuals have different breeding dispersal strategies following reproductive failure and that their apparent adult survival differences are at least partly explained by different breeding dispersal propensities. Our results also suggest that the recent decline of the study population reflects a multiple response to increased nest predation through decreased local recruitment and increased emigration.     

Effect of predation by skuas on breeding success of the Cape petrel Daption capense at Nelson Island, Antarctica

Breeding success of Cape petrels at Nelson Island (South Shetland Islands) in 1991/1992 averaged 29%. Predation by skuas accounted for 76% of all nesting failures. Direct attacks prevailed over opportunistic predation. Intensity of observer disturbance alone could not account for the inter-colony variation in predation rate, which tended to increase with colony size, presence of nearby breeding skuas and local availability of other prey species (penguins). Predation was significantly lower on nests that could be attacked only, or more easily, from the air and nests with close neighbours. Selective predation on nests in which relatively narrower eggs were laid was independent of nest-site characteristics and laying date, suggesting an effect of parental age/experience on breeding success through antipredator behaviour. Development of oil-spitting behaviour in growing chicks balanced the parental attendance and effort in nest defence, which declined as chick age increased. Increased predation on lighter but not smaller chicks may reflect the lower ability of such chicks to defend themselves by spitting stomach oil. Seasonal variations of predation rate on Cape petrel nests did not result from predator swamping by numbers of simultaneously active nests, but rather reflected temporal availability of alternative prey. Success of all phases of nesting declined with later laying date. Received: 15 December 1997 / Accepted: 11 April 1998     

Urban and coastal breeding lesser black-backed gulls (Larus fuscus) segregate by foraging habitat

Despite urbanization's general erosion of biodiversity, towns and cities provide novel opportunities for some species. During the 20th century, gulls (Laridae) colonized urban areas around the world where they flourished. At the same time, some coastal populations declined. The reasons for this difference are not fully understood, partly because little is known about any ecological differences between urban and non-urban gulls, such as their foraging ecology. Here we compare the movement ecology and habitat selection of Lesser Black-backed Gulls Larus fuscus graellsii breeding at two neighbouring colonies – one urban and one coastal – in north-west England. We used bird-borne GPS loggers to first compare colony-level movement behaviour and habitat selection and then investigated individual-level habitat use. We observed clear colony-level habitat segregation: urban breeders preferentially foraged in urban areas whereas coastal breeders foraged primarily in coastal habitats and avoided urban areas. Coastal breeders also had larger core and home-ranges than urban breeders, possibly due to differences in colony size. However, we also found inter-individual differences in habitat use, which may have important management implications. These findings suggest a link between nesting and foraging ecology, and thus management or environmental change altering food availability will impact gulls at the coastal and urban sites differently.     

Dispersal and recruitment during population growth in a colonial bird, the great cormorant Phalacrocorax carbo sinensis

While the factors influencing reproduction and survival in colonial populations are relatively well studied, factors involved in dispersal and settlement decisions are not well understood. The present study investigated exchanges of great cormorants Phalacrocorax carbo sinensis among six breeding colonies over a 13‐year period when the breeding population in Denmark increased from 2800 to 36 400 nests. We used a multistate capture‐recapture model that combined multisite resightings and recoveries to examine simultaneously recruitment, natal dispersal, breeding dispersal and annual survival of first‐year, immature and breeding great cormorants. Mean survival of first‐year birds (0.50±0.09, range=0.42–0.66 among colonies) was lower than survival of breeders (0.90±0.06, range=0.81–0.97). Mean survival of immature birds over the study period was 0.87±0.08. Dispersal from a colony increased with decreasing mean brood size in the colony in both first‐time and experienced breeders. The choice of the settlement colony in first‐time breeders was affected by conditions in the natal colony and in the colonies prospected during the pre‐breeding years. In particular, first‐time breeders recruited to colonies where they could expect better breeding success. Experienced breeders relied mainly on cues present early in the season and on their own breeding experience to choose a new breeding colony. Newly established colonies resulted mainly from the immigration of first‐time breeders originating from denser colonies. Dispersal was distance‐dependent and first‐time breeders dispersed longer distances than breeders. We suggest that the prospecting behaviour allows first‐time breeders to recruit in nearby as well as more distant potential breeding colonies. Dispersing breeders preferred to settle in neighbouring colonies likely to benefit from their experience with foraging areas. We discuss the importance of these movements for growth and expansion of the breeding population.     

Parental body size affects meal size in the chinstrap penguin (Pygoscelis antarctica)

Meal size of chinstrap penguins (Pygoscelis antarctica) was estimated through weighing adult breeders before and after food transfer during the crèche phase. Mean meal size delivered by each parent was 630.2 ± 178.3 g and was independent of breeding date, number of chicks and sex. Only body size as estimated by flipper length had a significant effect on meal size, suggesting an advantage of large body size which allows maximization of food carried per visit. Received: 22 September 1997 / Accepted: 16 December 1997     

Postmetamorphic dispersal and recruitment of first breeders in a Bufo calamita metapopulation

The migratory capacity of metamorphs and their later integration into local breeding populations was studied in a Bufo calamita metapopulation inhabiting the northern Rhineland, Germany (1991–1994). A new tagging method allowed unequivocal distinction among metamorphs (n = 1,709), which originated either from the spring-breeding cohort in 1991 and 1992 or from the summer-breeding cohort in 1991. The life history of batch-marked individuals was followed until adulthood, focusing on postmetamorphic dispersal, choice of breeding locality and timing of adult reproduction. Almost all metamorphs left the patches used for breeding within a few weeks after emergence. They were found at about the same density in all favourable microhabitats in a radius of up to 2 km around their native patch. This demonstrates that even small toadlets (6–11 mm snout-to-vent length) migrate distances of several hundred metres within a short time and that they significantly help prevent local extinction in sink populations (rescue effect). Recaptures of reproductive adults originating from tagged metamorphs were limited to males belonging to the 1997 spring and summer cohorts. The absence of recaptured females was probably due to their greater vagility and later sexual maturation. First-breeding males (2 years old) without exception joined the breeding cohort from which they originated: early breeders originated from spawn of the spring cohort, mid-summer breeders from spawn of the summer cohort. This result is in accordance with the hypothesis that the integration of first breeders to a specific breeding cohort is genetically controlled in central European natterjacks. The exchange of individuals between different cohorts in the same patch is considerably less than the migratory exchange between the same breeding cohorts among local populations, as predicted from the allozyme data. Received: 28 October1996 / Accepted: 4 June 1997     

Exploitation of inland-breeding Antarctic petrels by south polar skuas

During the austral summer of 1996/1997 we studied south polar skuas at Svarthamaren, Dronning Maud Land, Antarctica, where the world's largest known colony of Antarctic petrels is found. Our censuses suggested approximately 250 full-grown skuas and 140,000 breeding pairs of petrels were present. During their breeding season, skuas did not visit the open sea at least 200 km from the site; they relied entirely on prey caught and scavenged from the petrel colony. Because the site is so isolated, we asked whether the prey (petrels) had swamped the predators (skuas), or whether there was evidence that predator numbers were limited by the size of the prey population. Particularly at the end of the petrel incubation period, we found a close correspondence between the energy required by adult skuas and their chicks, ascertained from time budget studies, and the rate at which petrel eggs disappeared from the colony. This suggests that, in this closed system, the predator population was limited by the prey population, and that predator swamping was not an advantage that petrels gained by nesting in this remote location. Received: 12 April 1999 / Accepted: 30 June 1999     

The impact of maternal experience on post-weaning survival in an endangered arctic fox population

Behavioural differences in parental care can influence offspring survival through variation in e.g. antipredator behaviour and ability to provide food. In a broad range of species, offspring survival has been found to be higher for experienced females compared to inexperienced first-time breeders. The increase in offspring survival for experienced females has mainly been explained by improved experience in providing food. In this paper, we have studied post-weaning juvenile survival in relation to maternal experience in an endangered population of arctic foxes (Vulpes lagopus) in Fennoscandia. For cubs raised by inexperienced and experienced females, the survival rate was 0.42 (CI 95% ± 0.31) and 0.87 (CI 95% ± 0.08), respectively. There was no difference in body condition between the cubs and no observations of starvation. We suggest that the difference in survival was due to lack of experience to one of the most common predators, the golden eagle (Aquila chrysaetos). Golden eagles were mainly observed on dens with litters where the females were inexperienced first-time breeders. From a conservation perspective, it is therefore important to increase adult survival through actions to enlarge the proportion of experienced breeders.     

Stealing bivalves from common eiders: kleptoparasitism by glaucous gulls in spring

Here I report on glaucous gulls (Larus hyperboreus), an opportunistic, generalist predator, stealing bivalves from a diving duck, the common eider (Somateria mollissima). The study took place in spring, the pre-breeding period of the common eider, in an Arctic fjord (Adventfjorden) at western Spitsbergen, Svalbard. Eiders were abundant, their presence predictable, and they fed on large prey requiring surface handling—all factors facilitating food theft. Only adult glaucous gulls attended the eider flocks. The glaucous gulls brought stolen prey ashore. Amongst these the bivalve Mya neoovata (Myidae) was common. The probability that an eider flock was attended by glaucous gulls declined as the season progressed and increased with the foraging activity of the eiders. Eider flock size and the degree of aggregation within flocks were poor predictors of gull presence. However, eider flocks attended by a single gull were smaller than flocks attended by more than one gull. Common eiders are capital breeders which build up large energy reserves prior to breeding. Kleptoparasitism, therefore, may have a negative impact on eider energy acquisition in early spring. For the glaucous gull, kleptoparasitism may be important as few other food sources are available this time of the season.