Parental care of the Common Tern Sterna hirundo

Parental care activities of male and female Common Terns Sterna hirundo were recorded over two breeding seasons. Males and females exhibited distinct parental roles throughout a breeding bout. Courtship feeding by males was extensive prior to and during egg-laying, but declined with the onset of incubation. Females performed significantly more incubation behaviour than males although both sexes spent equal time attending at the nest site. During the chick stage, females spent significantly more time on the territory than did males. Chick feeding was largely the responsibility of the male; males fed chicks at a rate approximately three times higher than that of females. In addition, whereas females showed no trend in the size of fish delivered to chicks relative to chick age, the size of fish delivered by males increased with chick age. Courtship feeding activities and extensive chick feeding contributions by male Common Terns appear to outweigh parental contributions by females, contrary to predictions for a monogamous species.     

Effects of egg size, parental quality and hatch-date on growth and survival of Common Tern Sterna hirundo chicks

We examined the relative contributions of egg size, parental quality and hatch-date to growth and survival of second-hatched chicks (those chicks making the greatest contribution to differences in productivity among pairs) by exchanging clutches among nests of Common Terns Sterna hirundo matched for lay-date (range 13 May to 9 June). The mass of a second-laid egg in an exchanged clutch ranged from 17.70 to 23.80 g. Growth and survival were studied during three periods: early (days 0–3), middle (days 3–12) and late (days 12–25). Both egg mass and hatch-date were important predictors of hatchling mass (positive relationships), although there was no seasonal trend in egg mass. During the middle period, hatch-date was a significant predictor of mass gain and survival (inverse relationships). After controlling for hatch-date, other indices of parental quality made only small contributions to chick mass gain and survival. Our results suggest that although breeding early generally leads to greater overall survival of chicks, several important interactions among egg 'quality', parental quality and early laying may affect breeding success under specific conditions.     

Colony size and not nest density drives reproductive output in the Common Tern Sterna hirundo

Density is known to be an important factor in population size regulation. Several mechanisms of density limitation have been identified in colonial birds. We studied competition in Common Terns Sterna hirundo to assess whether the factor limiting reproductive output was competition for nest‐sites, which is dependent on local nest density, or density‐dependent competition for food resources, which is dependent on overall colony size using the same foraging area. We found strong associations of both colony size and nest density with reproductive output in five colonies of Common Terns in three different habitats (one marine, two freshwater). Based on detailed long‐term datasets of six separate sub‐colonies of the Banter See colony that differed in nest density, we found that reproductive success was not related to nest density but to overall colony size, possibly a result of resource depletion and food competition. We also found carry‐over effects of colony size during rearing on post‐fledging return rate. These results have important implications for the conservation management plans aimed at recovering declining populations of Common Terns.     

Feeding ecology of the Common Tern Sterna hirundo in a wintering area in southern Brazil

The wintering diet of Common Terns Sterna hirundo was studied by using 714 pellets collected on roosting sites at the mouth of the Lagoa dos Patos and on adjacent coastal beaches, in Rio Grande do Sul, southern Brazil, from March 1999 to February 2000. A total of 12 340 individual prey items of 35 different food types was found. Fish was the most important food type in the diet, constituting 32% by number and 93% by mass. Insects contributed 67% by number but only 3% by mass. The main food types were sciaenid fishes Paralonchurus brasiliensis, Micropogonias furnieri, Cynoscion guatucupa and Macrodon ancylodon. Several of these are important commercial species; fisheries potentially impact food availability to the terns, and terns may contribute significantly to the natural mortality of these fishes. Clupeiform fishes, the urophycid fish Urophycis brasiliensis and flying ants (Camponotus sp.) were also important. Species composition of the diet (food types), both by number and by mass, differed significantly between months. Prey sizes ranged in length from 12.7 mm to 217.4 mm. The average estimated total length of fish taken was 77.7 mm, but the mean differed significantly among prey species. The importance of demersal sciaenids to the diet of the Common Tern, a surface predator, may be explained by their association with aquatic predators, especially adult Bluefish Pomatomus saltatrix and Striped Weakfish Cynoscion guatucupa, and the Franciscana Dolphin Pontoporia blainvillei, which drive these fish to the surface. The occurrence of flying ants in the diet was related to offshore winds, which carried these insects out to sea. The occasional high availability of insects possibly changed the cost/benefit relationship of several food types, causing diet changes. The high number of prey species, the temporal variations in the composition of the diet and the wide range of prey sizes are evidence of the high dietary plasticity of the Common Tern, at wintering areas in southern Brazil.     

Intraspecific kleptoparasitism improves chick growth and reproductive output in Common Terns Sterna hirundo

Kleptoparasitism is a well‐known foraging strategy used opportunistically by many seabirds. Here, we investigated the effect of intraspecific kleptoparasitism on chick growth and reproductive output in Common Terns Sterna hirundo. Effects were compared between two groups comprising (1) individuals using kleptoparasitism during the chick‐rearing period (kleptoparasitic group, n = 18), and (2) individuals in pairs that never performed kleptoparasitism throughout the season (‘honest’ group, n = 21). The null models best described variation in mass at day 3 and the pre‐fledging mass, indicating no significant effect of the explanatory variables. However, the best models describing the linear growth rate (days 3–13) and peak mass included the parents' foraging strategy (kleptoparasitic vs. honest parents) as an explanatory variable. These two growth parameters were higher in chicks of kleptoparasitic parents. Kleptoparasitic foraging strategy was also associated with higher pre‐fledging survival, as the reproductive performance (i.e. number of fledglings) was significantly higher in the kleptoparasitic than in the honest group. We suggest that by stealing food (and consequently feeding offspring more frequently with high‐quality prey), kleptoparasitic parents are able to produce higher quality chicks with enhanced survival.     

Effect of GPS tagging on behaviour and marine distribution of breeding Arctic Terns Sterna paradisaea

Tracking tags have been used to map the distributions of a wide variety of avian species, but few studies have examined whether the use of these devices has impacts on the study animals that may bias the spatial data obtained. As Global Positioning System (GPS) tags small enough for deployment on terns (family: Laridae) have only recently become available, until now tracking of this group has been conducted by following unmanipulated individuals by boat, which offers a means of comparing distributions obtained from GPS‐tracking. We compared the utilization distributions (UDs) of breeding Arctic Terns Sterna paradisaea obtained by GPS‐tracking 10 individuals over 2 weeks, with UDs derived from contemporaneous visual boat tracks from 81 individuals. The 50% and 95% UDs of both methods had high similarity scores, indicating good agreement in the density distributions derived from the two methods. The footprints of the UDs of tagged birds were ~ 75–80% larger, which may reflect an effect of tagging on foraging range or the occasional inability to follow by boat individuals which roamed further from the colony. We also compared the nest attendance and chick provisioning rates of adults that were (1) fitted with a GPS tag and leg‐flag, (2) handled and marked with a leg‐flag but not tagged and (3) fitted with a leg‐flag in a previous year but unhandled in the year of the study. There was some evidence that birds fitted with both a GPS tag and leg‐flag spent slightly less time at the nest compared with unhandled birds and those fitted with a leg‐flag only. Both treatments where birds were fitted with a leg‐flag in the year of the study had similarly lower provisioning rates to those of unhandled control birds > 48 h after handling, suggesting that negative effects on provisioning are due to capture and handling or leg‐flag attachment rather than to GPS tag attachment/loading per se. Overall brood‐provisioning rate was compensated for by the increased effort by the unhandled partner. Our study suggests that despite slight effects of GPS‐tagging on behaviour, the estimates of marine density distribution obtained were very similar to those of unmanipulated birds.     

Variation in growth in Sandwich Tern chicks Sterna sandvicensis and the consequences for pre- and post-fledging mortality

Fitness consequences of variation in body mass growth and body condition were studied in a Sandwich Tern Sterna sandvicensis colony on Griend, Dutch Wadden Sea, during 1990–2000. Body mass increment during the linear growth phase predicted nestling survival probabilities accurately. Chicks growing less than 8 g per day had low survival probabilities until fledging, but within a range of 8–11 g per day growth only small effects on chick survival were observed. Effects of slow growth on survival became obvious after about 10 days after hatching. Slow growing chicks reached a much lower fledging mass, whereas slow growth had only small effects on structural size at fledging. Body condition of the chicks was highly variable and had strong effects on survival until fledging. However, body condition during the nestling stage did not influence post-fledging survival. Body condition at fledging had no effects on post-fledging survival and did not affect final mass or body size. It is argued that low fledging mass can be overcome soon after fledging, as parents take their fledglings closer to the foraging areas, thereby avoiding high rates of kleptoparasitism by Black-headed Gulls Larus ridibundus .     

Dispersal and climate change: a case study of the Arctic tern Sterna paradisaea

Dispersal is an important evolutionary process that can affect admixture of populations and cause rapid responses to changing climatic conditions due to gene flow from populations at different altitudes or latitudes already experiencing these conditions. We investigated long-term patterns of natal and breeding dispersal in a coastal seabird, the Arctic tern Sterna paradisaea , that experiences specific climatic conditions in the northern temperate and Arctic climate zones during breeding and different climatic conditions in the Antarctic during winter. Long natal and breeding dispersal distances were costly as shown by their effects on delayed breeding. Dispersal distances varied significantly among years, with natal dispersal showing a strong temporal increase during the last 70 years. Annual differences in dispersal distance could be accounted for by climate conditions in the breeding grounds and the winter quarters. Natal dispersal was related to climate conditions in both the year of hatching and the year of breeding, whereas breeding dispersal was only related to climate conditions in the second year of the dispersal event. Only the north Atlantic oscillation (NAO) index for winter showed a consistent temporal trend, suggesting that the temporal trend in natal dispersal distance must be caused by changes in the NAO (or associated phenomena). These findings indicate that dispersal can change rapidly in response to changing climate conditions.     

Overland movement and migration phenology in relation to breeding of Arctic Terns Sterna paradisaea

It is often implicitly assumed that seabirds migrate using marine environments, but this assumption is increasingly being challenged by electronic tracking data. The arrival and departure routes of Arctic Terns breeding on the North Sea coast of the United Kingdom (UK) are unknown but there has been speculation about the possibility of overland migration. Analysis of light-level geolocator data from birds breeding on the Farne Islands suggests that these birds arrived and left their North Sea colony overland via the Irish Sea, rather than taking coastal routes along the east coast of the UK and through the English Channel. In addition, some departing birds may enter the North Atlantic by crossing Ireland rather than through the Irish Sea. The direction of arrival in spring had a more southerly orientation than the direction of autumn departure. The geolocator data allow migration phenology in relation to breeding to be defined and indicated that the birds arrived around 15 days before the first eggs were laid in the colony. Departure timing may be determined by seasonal progression and not markedly influenced by breeding success. This study supports the idea that overland migration may be a more widespread and consistent strategy for seabirds than has been realized.     

The influence of age on the breeding biology and survival of the Arctic tern Sterna paradisaea

The paper describes the influence of the age of Arctic terns on the breeding biology and postfledging survival. Some of the effects of a "red-tide" on the terns are described.     

Effect of intraspecific interactions on seasonal decline in productivity of Common Terns Sterna hirundo

Capsule Kleptoparasitic activities of older chicks from earlier nests did not contribute to late reproductive declines.

Aims To determine whether intraspecific interactions, such as kleptoparasitism and aggression, were experienced more frequently by birds breeding late in the season as a result of exposure to breeders at a more advanced stage. If so, to investigate whether this was the cause of the observed seasonal decline in reproductive parameters observed at Bird Island, where nesting density is high and interactions are more probable.

Methods Plots were fenced within the colony, exploiting natural variability in distribution of early and peak breeders to create two treatments: plots with only late-laying terns and those with a mixture of early-, peak- and late-layers. Hatching success, productivity and the growth and survival of chicks were measured for all late-laying pairs. Intraspecific interactions, adult attendance and provisioning of chicks were recorded during 9600 minutes of nest observations made within two periods: a few days after hatching and one week later.

Results The frequency of intraspecific interactions was maintained by the kleptoparasitic activities of older chicks within the mixed-laying-date treatment and was significantly lower in plots containing only late breeders with chicks of similar ages (mean 11.0 days). The overall rate was rarely greater than two interactions per nest per hour and there was no corresponding reduction in the growth or survival of chicks from late nests or any change in the provisioning activities of late-breeding adults.

Conclusion Increased frequency of intraspecific interactions experienced by late breeders in the presence of early-breeding conspecifics resulted from the kleptoparasitic activities of older chicks but was not sufficient to contribute to the observed seasonal reproductive decline at this dense breeding colony.     

Conservation genetics of the Common Tern (Sterna hirundo) in the North Atlantic region; implications for the critically endangered population at Bermuda

Common Terns nesting at Bermuda are isolated by 1,000–4,000?km from other populations of the species around the North Atlantic Ocean. This population experienced a severe demographic bottleneck as a result of a hurricane in 2003 and was subsequently re-established by four males and four females. Using seven microsatellite loci, we compared the genetic diversity of the pre- and post-bottleneck populations, compared the genetic profile of the Bermuda population with those of other populations around the North Atlantic Ocean and mainland Europe, and assessed the potential contribution of immigration to genetic diversity. We found a loss of genetic diversity (number of alleles and heterozygosity) in the post-bottleneck Bermudian population (4.6–2.9 and 0.56–0.52, respectively). We also report significant differentiation among all sampled locations (global F_ST?=?0.16) with no evidence for immigration into Bermuda. Common Terns from the Azores were genetically more similar to those from mainland North America than to those from Bermuda or mainland Europe. Our results suggest that the critically endangered population in Bermuda is genetically distinct and requires continued and enhanced conservation priority.     

Mercury Concentrations in Common Tern Sterna hirundo and Slender-billed Gull Larus genei from the Shadegan Marshes of Iran,in North-western Corner of the Persian Gulf

We examined mercury levels in several tissues of Common Terns and Slender-billed Gulls collected from Shadegan Marshes of south-western Iran. In both species, total mercury content was highest in feathers followed by liver, kidney and muscle tissue. We found a significant correlation between mercury concentrations in kidney and breast feather (r?=?0.83, p?<?0.05), breast feather and liver (r?=?0.81, p?<?0.05) as well as liver and kidney (r?=?0.83, p?<?0.05). The contaminant levels in the feathers (11.53 and 15.32 μg/g in breast feather and tail feather, respectively) of Common Terns from Shadegan Marshes are higher than those reported for other tern species from elsewhere in the world, but feather mercury of Slender-billed Gull (6.61 and 5.35 μg/g in breast feather and tail feather, respectively) was similar to those reported for gull species worldwide. Mean values for mercury in the feather of two seabird species were higher than the levels known to cause adverse effects.     

Age‐dependent breeding dispersal and adult survival within a metapopulation of Common Terns Sterna hirundo

Dispersal is increasingly recognized as a process of fundamental importance in population dynamics and other aspects of biology. Concurrently, interest in age‐dependent effects on survival, including actuarial senescence, has increased, especially in studies of long‐lived seabirds. Nevertheless, datasets necessary for studying dispersal and age‐dependent effects are few, as these require simultaneous data collection at two or more sites over many years. We conducted a 22‐year capture‐mark‐recapture study of Common Terns Sterna hirundo at three breeding colonies 10–26 km apart in Buzzards Bay, Massachusetts, USA. All birds in the study were of known age (range 2–28 years, median 7 years, n = 3290) and 77% were of known sex. Estimates of adult recapture, survival and breeding dispersal rates were obtained for all age‐classes from 2 to 20 years. The model that acquired 100% of the QAIC_c (Akaike's Information Criterion adjusted for small sample size and overdispersion) weight in our analysis included age‐specificity in all parameters but no relationship with sex. Our study may be the first to demonstrate age‐specificity in recapture, survival and breeding dispersal rates simultaneously, using a single model. Annual rates of breeding dispersal ranged from <0.01 to 0.27, with a population‐weighted mean of 0.065; they decreased with increasing distance between colony sites and, unexpectedly, increased with age. Breeding dispersal did not increase consistently after years with predation on adults or after an attempt to displace birds from an oiled site. Survival rates did not vary among sites or years. Annual adult survival increased from 0.80 in 2‐year‐old birds to a maximum of approximately 0.88 around age 8 years and then declined to 0.76 at age 20 years, yielding strong evidence for actuarial senescence. The peak annual survival rate of 0.88 is at the low end of other estimates for Common Tern and in the lower part of the range recorded for other terns, but total numbers in the three colonies increased seven‐fold during the study. This was part of a slower increase in the regional population, with net immigration into the study colonies. Our results demonstrate the biological significance of breeding dispersal in local population dynamics and age‐related effects on survival and dispersal from a metapopulation of a long‐lived seabird.     

Post-fledging body mass increase in Common Terns Sterna hirundo: influence of age, sex and year

We studied the inter-year and inter-sex variation of the post-fledging body mass development of Common Terns Sterna hirundo in 2000 and 2001 at the Banter See colony, northern Germany. Here, post-fledglings can be identified and weighed remotely and automatically by a transponder system that makes use of automated balances installed at the colony. Individuals were sexed with PCR amplification methods. After fledging, young generally continued to increase their mass. However, in 2000, the young did not significantly increase their mass during the post-fledging period. In 2001, conditions were more favourable and body mass increased continuously. Further, in 2001, male post-fledglings were significantly heavier than female post-fledglings. Once having left the colony area (on average 18–23 days after fledging in 2000, and 14–16 days after fledging in 2001), post-fledgling body mass had still not reached adult body mass. The longer a juvenile stayed at the colony, the higher was its final body mass, which if acting as a threshold level may control departure time. Neither brood size nor hatching order affected post-fledging mass or period. In the unfavourable year 2000, when many individuals were found dead after fledging, fledging age but not fledging mass was found to be a predictor of post-fledging survival before departure: individuals fledging when older had a lower survival probability. Our results stress the importance of the post-fledging period for body mass increase and survival prior to departure. The variation in post-fledging mass growth between years and between the sexes is discussed with respect to parental effort and a possible selective provisioning of sons over daughters.