The influence of food supply on the breeding ecology of Kittiwakes Rissa tridactyla in Shetland

We measured the breeding performance, body condition, time budgets and foraging ranges of Kittiwakes Rissa tridactyla at Sumburgh Head, Shetland, in two years of contrasting food availability. Kittiwakes in Shetland generally feed their young almost entirely on sandeels, and fisheries data indicated that stocks of sandeels in Shetland waters were at least ten times higher in 1991 than in 1990. Fledging success of Kittiwakes was nil in 1990 and 68% of eggs laid in 1991, although clutch-size and hatching success were no different between years. Post-hatching foraging trips in 1991 were of comparable duration to those recorded at other colonies in conditions of good food supply (2–3 h), while trips recorded during incubation or post-hatching in 1990 were approximately three times longer on average than at corresponding stages of the breeding season in 1991. Radio-tracking data indicated that adults generally stayed within 5 km of the colony in 1991 but flew more than 40 km from the colony on each trip in 1990. Eggs were apparently not left unattended in either year, despite the fact that this required adults to incubate for periods in excess of 44 h in 1990. The extent to which adults were able to increase trip durations, foraging ranges and incubation shift lengths between years, while maintaining hatching success, indicates the degree to which Kittiwakes are normally buffered against adverse feeding conditions during incubation. Reduced nest attendance and lower body-condition of adults post-hatching in 1990, in conjunction with complete post-hatching breeding failure, indicate that adults were beyond the limits of their buffering capacity during chick-rearing in 1990.     

The effects of changes in food availability on the breeding ecology of great skuas Catharacta skua in Shetland

Great skuas on Foula, Shetland have responded to a decline in the availability of sandeels since the late 1970s by increasing the proportion of other items in their diets. This change is correlated with the annual recruitment of sandeels in Shetland waters. Since 1983 there has been a 10-fold increase in predation by great skuas upon other seabirds, as Furness & Hislop (1981) suggested might occur in response to a low availability of sandeels. Changes in diet have been accompanied by a 50% reduction in adult territorial attendance as adults increased their foraging effort, such that between 1987 and 1989 breeding adults were probably working as hard as they were able to. Despite this, breeding success was less than 40% in 1987 and less than 15% in 1988 and 1989. The major cause of breeding failure was predation of unguarded chicks by adults from neighbouring territories. The willingness of adults to expose their chicks to high predation risk is probably maintained because of a positive correlation between chick pre-fledging growth and post-fledging survival, which is expressed up to the age of two years and which will place a strong pressure upon adults to feed their chicks as well as possible. The high expenditure of effort by adults in 1987 and 1988 did not affect the weights of those birds incubating eggs in 1988 and 1989, but there was a slight (3%) decrease in egg size between the late 1970s and the late 1980s. Changes in the age structure of the breeding population and the absence in 1989 of 28% of adults colour-ringed during incubation in 1988 suggest an increase in the rate of egress since the 1970s. These changes probably represent an increase in the long-term costs of reproduction to adults at this colony.     

Agonistic display and spacing in the black-headed gull, Larus ridibundus

Black-headed gulls nesting on islands in a marsh habitat had a higher breeding success than those nesting on the periphery of the colony. Island birds maintained close inter-nest distances whereas peripheral nests were spaced further apart as predation of eggs increased. These changes in spacing corresponded with changes in the response distances of agonistic displays. There was no change during incubation for island birds but peripheral birds displayed to intruders at much greater distances than during pair formation. Island birds were more likely to attack and display more frequently to intruders than peripheral birds. However, in one season the peripheral birds were more aggressive during pair formation because of an increase in intrusions by unpaired females.     

Seabirds and sandeels: the conflict between exploitation and conservation in the northern North Sea

Sandeels, especiallyAmmodytes marinus are a major component of the diet of many predatory fish, seabirds and seals. The industrial fishery for sandeels is now the largest of the North Sea fisheries. A sandeel fishery in the Shetland area began in 1974 but has recently declined. This change was accompanied by dramatic declines in the breeding success of certain seabirds, particularly Arctic terns and kittiwakes. Current information on seabirds and sandeels in the Shetland area is reviewed and areas where further research is needed, highlighted. The Shetland problem illustrates the difficulties of reconciling conservation and exploitation when fundamental ecological and behavioural knowledge is lacking, and also the need to obtain further information on the ecological impact of industrial fisheries.     

Population trends in bird species at Dungeness, Kent

Most wintering wildfowl species increased in numbers on the Dungeness Reserve during the period 1969-87, particularly teal, gadwall and wigeon. Wigeon, teal and gadwall showed the fastest rates of increase. An exponential model provided a better fit to the data for smew and gadwall than a linear one. Populations were more or less as variable in number in the early years of management (1969-77) as in subsequent years (1978-86).
Of wintering and passage wader species, turnstone, redshank, black-tailed godwit and common sandpiper increased. Snipe showed a decline in abundance in spring. Numbers of black terns, the only non-wading migrant analysed, remained variable over the period.
Breeding tufted duck and great-crested grebe increased in numbers during the period 1969-86 whereas mallard declined. Black-headed gull numbers increased, as did exponentially-breeding common and sandwich terns. Of four passerine species analysed, only breeding reed warblers increased during the period 1968-86, whereas numbers of breeding sedge warbler and whitethroat remained stable. Wheatear numbers declined in early years but have increased since 1983.
Increases in species richness were documented in wintering and passage, and breeding species.     

Is the additional effort of renesting linked to a hormonal change in the common tern?

The production of and care for a replacement clutch can bear costs in terms of future reproduction or survival. However, renesting is quite common among seabirds and can contribute considerably to individual fitness. Prolactin and corticosterone are two hormones involved in the mediation of breeding behavior and, as they are linked to body condition or effort, it is of interest if these hormone values change during a second demanding breeding phase within a year. We compared baseline prolactin and corticosterone between the first and the renesting attempt in common terns (Sterna hirundo) on individual level. Therefore, in addition to control birds, 37 breeders were sampled during incubation of their first and their replacement clutch in 2008 and 2009. Blood samples were taken non-invasively by blood-sucking bugs. Prolactin level was lower during the renesting period, especially in birds which abandoned their clutch afterwards, whereas corticosterone did not change. Excluding the deserting birds, the reduced prolactin level was not linked to minor success, but could be related to seasonal processes. The control group of late laying common terns showed comparably low prolactin values, but increased corticosterone concentrations. Renesting individuals exhibited higher prolactin during incubation of their first clutch than non-renesting birds, probably indicating their higher quality. The fact that terns still have relatively high prolactin and low corticosterone values during renesting might confirm their higher quality and suggests that they are able to meet the costs of a second demanding breeding period without being considerably stressed.     

Predicting reproductive success from hormone concentrations in the common tern (<Emphasis Type="Italic">Sterna hirundo</Emphasis>) while considering food abundance

In birds, reproductive success is mainly a function of skill or environmental conditions, but it can also be linked to hormone concentrations due to their effect on behavior and individual decisions made during reproduction. For example, a high prolactin concentration is required to express parental behaviors such as incubation or guarding and feeding the young. Corticosterone level, on the other hand, is related to energy allocation or stress and foraging or provisioning effort. In this study, we measured individual baseline prolactin and corticosterone between 2006 and 2012 in breeding common terns (Sterna hirundo) using blood-sucking bugs. Reproductive parameters as well as prey abundance on a local and a wider scale were also determined during this period. Baseline prolactin and corticosterone varied significantly between years, as did breeding success. At the individual level, prolactin was positively and corticosterone was negatively linked to herring and sprat abundance. At the population level, we also found a negative link between corticosterone and prey abundance, probably reflecting overall foraging conditions. High prolactin during incubation was mainly predictive of increased hatching success, potentially by supporting more constant incubation and nest-guarding behavior. It was also positively linked to a lesser extent with fledging success, which could indicate a high feeding rate of young. Corticosterone concentration was positively related to high breeding success, which may be due to increased foraging activity and feeding of young. In general, our study shows that baseline prolactin and corticosterone levels during incubation can predict reproductive success, despite the presence of an interval between sampling and hatching or fledging of young.     

Diet of Black-legged Kittiwakes (Rissa tridactyla) feeding chicks at two Irish colonies highlights the importance of clupeids

Capsule We examined regurgitates from Black-legged Kittiwakes during the early chick-rearing period over two breeding seasons at two colonies in Ireland where diet has not been studied previously. Clupeids were the dominant food source at both colonies, which contrasts with other studies throughout the British Isles that suggest Kittiwakes feed mainly on sandeels during this period. Our study is limited by sample size and restricted to the early breeding season, but suggests a link between reproductive success and dietary composition which should be investigated further.     

Atlantische Papagei‐taucher in Not

Puffins, climate change and fisheries Long term changes in north Atlantic weather leading to climate changes have measurable influences on marine eco systems. Seabirds have to cope with changes in the food web, driven by rising SST and other abiotic factors. The Atlantic Puffin is a good example for this. We can study its breeding success and colony size as measurements for changes in the food resources, mainly sandeels or herring over a wide area from the southern boreal to the arctic waters and from the east to west coast colonies of the Atlantic ocean. The findings show that population changes and breeding success in several seabird species like the Puffin and Common Guillemots or Black‐legged Kittiwakes as well show strong correlations to declining sandeels as a consequence of intensive industrial fishing, changes in SST and plankton, especially the most abundant copepod Calanus finmarchicus, development.     

Aggression by adult South American Terns toward conspecific chicks

ABSTRACT Infanticide and nonfatal aggression by adults toward unfamiliar chicks have been widely reported in colonial birds, and can be an important cause of chick mortality. We studied intraspecific aggression by adults toward chicks at a South American Tern (Sterna hirundinacea) colony in Patagonia, Argentina, during 2005 to characterize this behavior, evaluate its relationship with nesting density, chick age and microhabitat characteristics, and assess its effect on breeding success. Of 111 chicks in the study area, 45% were attacked at least once. Chicks older than 9 d posthatching were more likely to be attacked than younger chicks, and unattended chicks were more likely to be attacked than guarded chicks (88 vs. 12%). Chicks were also attacked more often when in their own territories (76% of cases), but were less likely to be attacked in territories with more vegetation cover and high‐quality shelters (i.e., vegetation with characteristics that prevented adults from reaching chicks). The number of aggression events was not related to nest density. At least 8% of the chicks in our study area died as a result of adult intraspecific aggression. Our results indicate that intraspecific aggression by adult South American Terns toward chicks is relatively common in the Punta Loma colony and should not be underestimated as a factor affecting their breeding success.     

Effects of individual life-history traits and weather on reproductive output of Black-headed Gulls Larus ridibundus breeding in the Wadden Sea, 1991–97

Capsule Annual breeding success was relatively constant and is shown to be related to clutch size and growth rate and to be positively affected by rain during egg-laying and advanced chick-rearing phase.

Aims To provide the first long-term data on breeding success of Black-headed Gulls in the Wadden Sea, to analyse its intrinsic and environmental co-variates, and to re-assess the significance of the severe winter 1995/96 on reproduction.

MethodsIn a relatively small colony at the Wadden Sea coast, clutches were selected randomly and enclosed to determine clutch size, egg biometrics, hatching and breeding success, and chick development in 1991 and from 1994 to 1997. Weather data (temperature, rain, wind) were related to life-history traits (clutch size, egg volume, chick development) and reproductive success.

Results Mean annual breeding success was 0.7 fledglings per pair. In 1996, breeding success tended to be lower and chick growth rate was significantly lower. Hatching success was lower in small clutches, chick mortality increased with decreasing chick growth rate. Rain during the egg-laying phase increased clutch size and decreased clutch predation. Rain during the early post-natal phase impaired chick growth. Rain during the phase of linear growth affected chick growth and fledging success positively and brood predation negatively.

Conclusions Breeding success of Black-headed Gulls breeding in the Wadden Sea is relatively constant between years, probably due to the use of terrestrial and marine feeding habitats. Rain may increase the availability of intertidal and terrestrial prey and thus may affect time budgets and food provisioning of parents positively. Lower breeding success in 1996 might have been caused by a relatively dry breeding season and possibly by the preceding severe winter.     

Common Guillemot Uria aalge parents adjust provisioning rates to compensate for low food quality

The quantity and quality of food available within the foraging area set important constraints for chick‐rearing birds, but responses to low quality are not well understood. This study explored the potential for parent birds to adjust quantity (feeding rate) and quality (energy content) in chick provisioning, by studying Common Guillemots Uria aalge on Stora Karlsö, Baltic Sea, predominantly utilizing Sprat Sprattus sprattus, during conditions of high food quantity but reduced food quality. Quality is central to reproductive success in this single‐prey loader. From the chick's perspective, provisioning rates should be increased to compensate for low food quality and to fulfil its growing needs with increasing age. However, the high energy cost of flying in Guillemots makes it important for parent birds to minimize commutes to feeding areas. Provisioning parameters were recorded during three dawn‐to‐dusk watches each breeding season from 2005 to 2013, when clupeids, presumably Sprat, constituted 98% of chick diet. Generalized additive mixed models showed that both feeding rate and size of clupeids (a proxy for energy content) varied between years and changed non‐linearly with chick age, but that there was no change within breeding seasons. Chick age and year explained 36% of the variation in feeding rate but only 2% of the variation in the size of clupeids in chick diets. We conclude that parent birds tried to adjust both feeding rate and prey size, but were less successful with the latter. A strong negative correlation was found between annual feeding rates and size of clupeids, evaluated as the differences relative to the baseline year, and adjusted for the effects of chick age. Although the differences between years were small, the relationship indicates a compensation mechanism that does not seem to impact adult survival, and by which increased feeding rates can partly counteract reduced chick energy intake when food quality is low.     

Little tern breeding success in artificial and natural habitats: modelling population growth under uncertain vital rates

As a consequence of habitat loss, breeding in man-made habitats has become increasingly common for many coastal breeding bird species. While artificial sites provide valuable substitutes, they may also be more attractive, and importantly, differ in quality from natural sites. Therefore, information on habitat specific breeding success and their potential for supporting stable populations are needed. We compared little tern (Sternula albifrons) breeding success (nest and hatching success) between natural habitat (sandy beaches) and artificial port habitat at Bothnian Bay, Finland from 2006 to 2011. We further reviewed published estimates on pre-fledging and adult survival for little terns and least terns (Sternula antillarum), and used these ranges to estimate plausible parameter spaces for population growth rates given our estimates of breeding success. Nest success was among the highest reported for little terns in the artificial habitat (82 %) while being lower in the natural habitat (58 %). This difference may have resulted from differences in colony sizes and levels of disturbance. Hatching success did not differ significantly, but the percentage of successful nests containing unhatched eggs was twice as high in the natural habitat. The parameter spaces for population growth rates indicated that the artificial habitat has good potential to sustain stable populations (66 % positive growth rate) while for the natural habitat this potential was lower (37 % positive growth rate). While our results suggest that artificial habitats can be very productive breeding sites for habitat deprived tern populations, management should concentrate on improving both habitats with emphasis on natural sites.     

Wie richtet eine Flußseeschwalbenkolonie (Sterna hirundo) ihr Abwehrverhalten auf den Feinddruck durch Silbermöwen (Larus argentatus) ein?

How a Common Tern (Sterna hirundo) Colony Defends itself against Herring Gulls (Larus argentatus) The subject of this study is the anti-predator behaviour of a small common tern colony near a large herring gull colony on the island of Mellum, West Germany (Fig. 1). In 1980 the number of gulls crossing this tern colony increased during the terns' chick-stage (Fig. 4). Observed predation of tern chicks was independent of tide and time of day (3., Fig. 5). The frequency of tern reactions corresponded to the number of herring gull crossings (Fig. 5, Table 1). The terns' responses increased between morning and evening (Fig. 8). Tern up-flights and attacks increased absolutely and as a percentage, with the advance of the breeding season (Fig. 3, 4). They were positively correlated with the observed chick predation and the number of pairs with chicks, most markedly with chicks older than 5 days (Figs. 3, 4; Table 1). This increased defence was maintained by fewer pairs as, by then, many had lost their own broods (Fig. 4). As the breeding season progressed, herring gulls increasingly became the main cause of tern up-flights and the object of the attacks (Figs. 9–11). The up-flights of the whole colony, which occurred frequently and spontaneously during incubation, were observed only rarely after hatching and were almost exclusively a response to herring gulls (Figs. 10, 12). The lower herring gulls flew over the colony, the more frequently common terns flew up or attacked and the more individuals were involved in these responses (Figs. 6, 13, 14). During the breeding period, communal up-flights and attacks by terns increased as a percentage (Figs. 12, 13, 15–17). Group-attacks effected changes in the gulls' flying-routes more often than did individual attacks (Fig. 18). Despite the defence behaviour and its adaptation to the predation pressure, herring gulls often succeeded in robbing chicks. This is why the breeding success of the common tern was poor (< 0.4 chicks/nest). Possible reasons for this are discussed.     

The diet of Common Guillemot Uria aalge chicks provides evidence of changing prey communities in the North Sea

Local differences in feeding conditions have been suggested as a cause of regional variation in seabird demography but multi‐colony comparisons of diet are rare. In UK waters the main fish eaten by seabirds during the breeding season belong to three families: Ammodytidae, Clupeidae and Gadidae. Climate change and fishing are affecting these fish stocks and so probably impact on predators such as seabirds. We used standardized observations of prey brought in for chicks to make the first integrated assessment of the diet of Common Guillemot Uria aalge chicks at a UK scale. Chick diet varied markedly among the 23 colonies sampled between 2006 and 2011. Sandeels (Ammodytidae), probably Lesser Sandeels Ammodytes marinus, were the commonest prey. Their contribution to the diet varied both latitudinally and among marine regions, with the proportion significantly higher for a given latitude on the west coast compared to the east. The non‐sandeel component of the diet showed latitudinal changes, with small clupeids, probably Sprats Sprattus sprattus, predominant at southern colonies whereas juvenile gadids were the main alternative to sandeels in the north. Comparison of our Guillemot chick diet with data collected 15–30 years earlier suggests that the proportion of sandeels in the diet has decreased at colonies bordering the North Sea. No significant change was apparent in Atlantic colonies but historical data were limited. The early years of our study coincided with a population explosion of Snake Pipefish Entelurus aequoreus in the Northeast Atlantic and North Sea. Pipefish were recorded in Guillemot chick diet at several northern and northwestern colonies in 2006 and 2007 but have been absent since 2009. Spatial and temporal variation in chick diet accorded broadly with patterns expected as a result of rising sea temperatures and impacts of fishing. Guillemot chick diet could potentially be a useful indicator of changes in the distribution and abundance of forage fish.     

Sex roles, parental experience and reproductive success of eastern kingbirds, Tyrannus tyrannus

We quantified parental behaviour of eastern kingbirds during the incubation and nestling periods to determine parental roles, and to examine the impact of previous breeding experience (defined as having bred on the territory in the past) on behaviour and reproductive success. Females performed all incubation, while males spent more than 60% of their time in vigilant or nest guarding behaviour during incubation. Parental roles were not defined as sharply during the nestling period. Females spent more time vigilant, but males provisioned young at only 54% of the rate of females. Vigilance and nest watching were still primarily male duties. Male and female behaviour did not vary with the pair's combination of experience (e.g. experienced-experienced versus inexperienced-inexperienced in previous-current breeding season, respectively) during either phase of reproduction, but experienced males were more vigilant during incubation and fed young relatively more than inexperienced males. Experienced females were also more efficient foragers. Although behaviour did not differ among the four combinations of pair experience, inexperienced pairs none the less lost the most young to starvation and predation. Consequently, inexperienced pairs fledged one less nestling per nesting attempt than did pairs with at least one experienced breeder. Our results suggest that having at least one experienced breeder substantially improved a pair's reproductive success. We propose that female site fidelity is a safeguard to avoid the lower breeding success a female would incur if she were to move to a new territory and breed with an inexperienced male. Copyright 1999 The Association for the Study of Animal Behaviour.     

Improved estimates of digestion correction factors and passage rates for harbor seal (Phoca vitulina) prey in the northeast Atlantic

Diet composition in pinnipeds is widely estimated using hard prey remains recovered from feces. To estimate the size and number of prey represented in fecal samples accurately, digestion correction factors (DCFs) must be applied to measurements and counts of fish otoliths and cephalopod beaks. In this study, 101 whole prey feeding trials were conducted with six harbor seals (Phoca vitulina) and 18 prey species. We derived species‐ and grade‐specific estimates of digestion coefficients (DCs) and species‐specific recovery rates (RRs) to account for partial and complete digestion, respectively. Greater than 98% of otoliths were passed within three days of consumption. RRs were smallest for Atlantic salmon smolts (RR = 0.306, SE = 0.031) and increasingly larger for sandeels (RR = 0.494, SE = 0.017), flatfish (RR = 0.789, SE = 0.033), and large gadoids (RR = 0.944, SE = 0.034). Species‐specific otolith width DCs were smallest for Trisopterus species (DC = 1.14, SE = 0.015) and increasingly larger for flatfish (DC = 1.27, SE = 0.045), large gadoids (DC = 1.32, SE = 0.067) and sandeels (DC = 1.57, SE = 0.035). RRs were similar to those from gray seals (Halichoerus grypus), but harbor seal species‐ and grade‐specific DCs were generally smaller. Differences in partial and complete digestion rates among prey species and between seal species highlight the importance of applying DCFs when reconstructing diet.     

THE KLEPTOPARASITIC BEHAVIOUR OF THE ARCTIC SKUA STERCORARIUS PARASITICUS WITH THREE SPECIES OF TERN

The behaviour of Arctic Skuas when stealing fish from three species of tern was studied at the estuary of the River Ythan, Aberdeenshire. The skuas' attacks were made by flying low over the water at high speed and by approaching the terns from behind and below. The adaptiveness of this flight path is discussed. The skuas attacked singly and in groups of up to five. Group attacks were more successful than single bird attacks, but the return per bird was greatest when the skuas attacked in pairs. The success of attacks also depended upon the species of tern being attacked. The duration of attacks varied with the number of skuas in the attack, the species of tern being attacked and the outcome of the attack. Attacks were directed more frequently against Sandwich Terns than against Common Terns, correlating with the success of the attacks against these two species.     

Species interactions in a mixed colony of common terns (Sterna hirundo) and black skimmers (Rynchops niger)

Black skimmers (Rynchops niger) frequently nest among common terns (Sterna hirundo) on coastal islands from North Carolina to New York. Behavioural interactions among terns, skimmers, and several potential avian predator species were quantified during the 1973 breeding season at Fisherman Island, Virginia. Common terns were dominant to skimmers, chasing and dive-attacking neighbouring pairs, while skimmers rarely chased terns. Despite agonistic interactions, nesting success of both species was no different whether nearest neighbours were conspecifics or heterospecifics. Common terns chased away potential avian predators much more frequently than did skimmers. Black skimmers, then, appear to derive a ‘protective cover’ benefit from nesting among the terns. Neither nest-site limitation nor food ‘information-sharing’ appear to account for the species associations.     

Compared distribution within a disturbed fishpond ecosystem of breeding ducks and bird species indicators of habitat quality

The distribution of breeding ducks versus three other bird species (Mute Swan Cygnus olor, Black-headed Gull Larus ridibundus and Purple Heron Ardea purpurea) among waterbodies was investigated in the Dombes area, Eastern France, where breeding duck populations have undergone a severe decline following a large-scale transformation of meadow habitat into cropland. Higher Pochard Aythya ferina, Tufted Duck Aythya fuligula, Red-crested Pochard Netta rufina and Gadwall Anas strepera pair densities were recorded in fishponds where Black-headed Gulls were nesting. In these ponds however, nesting success (assessed by the number of broods divided by the number of pairs) was not significantly higher. Similarly, Pochard pair density was higher in ponds with a Purple Heron colony, but brood densities were not. We hypothesise that, in the study region, clutch concentration in the most attractive areas could compensate for the anti-predation effect of gull colonies or overwater nesting in shore vegetation. We could not confirm the expected negative impact of Mute Swan aggressive behaviour on duck distribution. Even though Mallard Anas platyrhynchos pair density was lower in ponds with breeding swans, we did not observe difference for Mallard broods. Moreover, Red-crested Pochard pair density and nesting success were higher in ponds where swans were breeding, probably as a consequence of shared habitat preferences.