The incidence of the tick parasite Ixodes uriae in Kittiwake Rissa tridactyla colonies in relation to the age of the colony, and a mechanism of infecting new colonies

Kittiwakes Rissa tridactyla (683 chicks and 64 adults) were examined for ectoparasites in three breeding areas: Isle of May (Firth of Forth, Scotland; eight colonies of approximately known age of occupation on natural sites), North Shields and Gateshead (Tyne and Wear, England; well established colonies of known age on buildings). All the 208 ticks collected were Ixodes uriae. On the Isle of May, a colony which had only been established for 7 years was already infested by ticks. Tick incidence increased significantly with the length of colonization of the area during the first 30 years. Age of colonization did not have an effect after 30 years. In Gateshead only a single adult among the 127 birds examined had a tick. No ticks were found on the 106 chicks born at North Shields checked in 1987 or in 1988. However, among nine visiting newly-fledged chicks from neighbouring colonies caught at the North Shields colony, six had either ticks or scars of previous infestation. Some visiting chicks remained for several days in the host colony and were even fed by the owners of visited nests. Ixodes uriae may thus spread from one colony to another during visits of newly-fledged infested chicks to neighbouring colonies. The absence of any infestation in old colonies (38 years in North Shields and 17 years in Gateshead) on buildings that are regularly visited by newly fledged infested chicks, suggests that buildings may lack the rock refuges necessary for tick survival from one year to the next. In this respect Kittiwake colonies on buildings are rather atypical.     

Individual Vocal Recognition in the Kittiwake Gull,Rissa tridactyla (L.)

Breeding Kittiwake Gulls recognized their partners by voice alone. The voices of all individuals in the population showed consistent differences in their tonal and temporal structures. The role of calling behaviour in a cliff-nesting context is discussed.     

Parental attendance and squatting in the Kittiwake Rissa tridactyla during the rearing period

Parental attendance was studied in 1991 in the Cap Sizun Kittiwake colonies (Brittany, France). After a period of continuous guarding lasting on average 22 days, parents left their chicks unattended. Thereafter, parental attendance decreased regularly until fledging. The chick age when first left alone was on average 3 days lower for large broods than for single-chick broods. Moreover, whatever the brood size, chicks from late nests were younger when left unattended. Parental age affected the initiation of first absence. Younger parents reduced their attendance sooner than older parents. About 80% of the nests with chicks were visited by other adults at least once during the absence of the parents, and 50% were visited in the 3 days following the first absence of the breeders. These squatters were mainly failed breeders and prebreeders looking for a future breeding site. The results are discussed in terms of costs and benefits of chick neglect and comparisons were made with data from other studies in North Atlantic and Alaskan colonies. This reflected the flexibility of adult behaviour in relation to brood size and food availability.     

The energy economy of the arctic-breeding Kittiwake (Rissa tridactyla): a review

The present paper reviews recent studies on changes in body mass, body composition and rates of energy expenditure during the breeding season in the black-legged Kittiwake (Rissa tridactyla) on Svalbard (79 degrees N). The main characteristic of the energy budget is a pronounced decrease in body mass as well as basal metabolic rate (BMR) after the eggs have hatched. While most internal organs lose mass in direct proportion to the general decrease in body mass, the liver and kidney masses decrease to a disproportionately greater extent. Since both the liver and the kidney have high intrinsic metabolic rates, these results support an earlier notion that the reduction in body mass is an adaptation to reduce maintenance costs. Alternatively, the reduced BMR is due to a decrease in energy uptake from the gastrointestinal tract, thereby ensuring that undigested food is ready to be regurgitated to the chicks. At the end of the chick-rearing period, the field metabolic rate (FMR) reaches its highest level, probably due to an increased workload associated with chick feeding. This occurs at a time of low body mass and BMR. A pronounced increase in the metabolic scope (FMR/BMR) during the latter part of the chick-rearing period demonstrates that BMR and FMR may change independently of each other and that the ratio FMR/BMR may not be a good measure of energy stress.     

The influence of pair-status on the breeding behaviour of the Kittiwake Rissa tridactyla before egg-laying

The breeding behaviour of individually marked Kittiwakes Rissa tridactyla that retained mates from the previous year (SAME) was compared, over the period from pair-formation to egg-laying, with those that changed mates (CHANGE). Position of nest-site in colony and breeding experience did not differ in the two groups studied. Hirds in CHANGE pairs attended the nest-site more frequently and as a result probably had less 'off-duty' time during which to forage. Rates of greeting were higher in CHANGE pairs. Differences between the two groups were usually greatest during the first two weeks after pair-formation an decreased toward egg-laying. CHANGE pairs copulated over a longer period before egg- laying, but stopped copulating sooner than SAME pairs. Copulation rates were higher in SAME pairs immediately before egg-laying. The causation of the behavioural differences between the two groups is discussed in terms of mate familiarity. It is suggested that the recorded differences explain the lower reproductive success in CHANGE pairs reported by other workers.     

Breeding ecology of the Fulmar Fulmarus glacialis and the Kittiwake Rissa tridactyla in high-arctic northeastern Greenland, 1993

The breeding ecology of the Fulmar Fulmarus glacialis and the Kittiwake Rissa tridactyla in the high Arctic was studied in relation to the occurrence of the northeast water polynya in northeasternmost Greenland (80̀N). Mean laying dates were 31 May in the Fulmar and 18 June in the Kittiwake; the total nesting season for the Fulmar just matched the time window of the polynya opening period. Fulmar colony attendance fluctuated within a period of 11.6 days because of variation in nonbreeding prospectors but showed no clear diurnal variation. Fulmar incubation shifts, on average, lasted 6.1 days (range 1–13 days), which is significantly longer than elsewhere, and the average chick-guard period of 10.9 days (range 1–17 days) was significantly shorter than in other studies. Egg neglect occurred in 18% of Fulmar nests or 0.7% of nests per day. Overall breeding success (chicks fledged per egg laid) was 0.56 in the Fulmar and 0.67 in the Kittiwake; the latter produced 1.4 young per active nest or 1.2 per completed nest. Mean Kittiwake clutch size was 2.03; larger clutches were laid early. Nest site characteristics (presumably reflecting nest predation risk) and breeding behaviour affected breeding success. in the Fulmar, hatching success was negatively correlated with laying date and the proportion of egg neglect, while overall breeding success was correlated negatively with distance to nearest neighbouring site and positively with the length of the chick-guard period. Kittiwake breeding success was negatively correlated with laying date. Using seabirds as indicators of marine food supply, breeding success in both species suggested moderate to good food supply in the northeast water polynya in 1993, although at least in the Fulmar the high reproductive output appeared partly maintained by behavioural buffering; long incubation shifts, egg neglect and short chick-guard periods were symptoms of foraging constraints.     

Maternal Androgens Increase Sibling Aggression,Dominance, and Competitive Ability in the Siblicidal Black-Legged Kittiwake (Rissa tridactyla)

Animals and plants routinely produce more offspring than they can afford to rear. Mothers can favour certain young by conferring on them competitive advantages such as a leading position in the birth sequence, more resources or hormones. Avian mothers create hatching asynchrony within a clutch and at the same time bestow the eggs with different concentrations of androgens that may enhance or counteract the competitive advantage experienced by early-hatching “core” young. In siblicidal birds, core young assume a dominant social position in the nest due to their size advantage and when threatened with starvation fatally attack subdominant later-hatching “marginal” young. A role for maternal androgens in siblicidal aggression has frequently been suggested but never tested. We studied this in the facultatively siblicidal black-headed kittiwake. We found that marginal eggs contain higher instead of lower concentrations of androgens than core eggs. Surprisingly, exposure to experimentally elevated yolk androgens increased sibling aggression and dominance, even though in nature marginal eggs never produce dominant chicks. We propose the “adoption facilitation hypothesis” to explain this paradox. This cliff-nesting colonial species has a high adoption rate: ejected marginal kittiwake chicks frequently fall into other nests containing chicks of similar or smaller size and exposure to yolk androgens might help them integrate themselves into a foster nest.     

Reproductive success of the Kittiwake Rissa tridactyla: the roles of clutch size, chick growth rates and parental quality

Kittiwake growth rates and breeding success are examined in relation to survival between fledging and breeding and to adult survival rates. High chick growth rates lead to increased survival after fledging. Broods of three (the maximum brood size) did not suffer lower fledging success than broods of two and clutches of three fledged appreciably more chicks per pair than did clutches of two or one. On average, the a- and b -chicks in broods of three grew at a faster rate and had a higher survival before breeding than those from smaller broods. Chicks from broods of two with experienced female parents grew at a faster rate than those of inexperienced female parents. Female parents which laid three egg clutches had a higher survival rate than those which laid clutches of two or one. We contend that three egg clutches were laid by higher quality individuals. We believe that clutch size indicates the condition of the Kittiwakes forming the pair. This condition probably has a genetical component, but is modified by environmental factors.     

Changes in body condition in breeding kittiwakes Rissa tridactyla

We investigated the seasonal pattern of changes in body condition of breeding black‐legged kittiwakes Rissa tridactyla in Svalbard (79°N) to evaluate whether changes in body condition were a consequence of the energetic demands of breeding (the reproductive stress hypothesis) or of voluntary anorexia to attain lower flight costs during chick rearing (the programmed anorexia hypothesis). Adult body condition was recorded from early egg laying to fledging and was examined in relation to date (relative to hatching), sex, parental time‐budget, brood size and reproductive success. To distinguish between the two hypotheses we evaluate whether the reduction in body condition occurred during or ahead of the energetically most demanding part of the chick‐rearing period. We combine our results on changes in body condition and time‐budget with published information on field metabolic rate (FMR) and chick energy requirements from studies in the same colony.
Our calculations of adult energy requirements and energy intakes indicate that the first part of the chick‐rearing period was energetically the most demanding period, because adult energy requirement per hour spent off the nest was highest in this period, and adults were time constrained because of the need for 24‐h brooding of the chicks.
During the incubation period female body condition increased slightly, but significantly, while male body condition was stable. During the first part of the chick‐rearing period, female and male body condition decreased by 14.8% and 8.4%, respectively. During the second part of the chick‐rearing period, both male and female body condition remained stable. The reductions in body condition occurred during the phase which was suggested to be the energetically most demanding part of the chick‐rearing period, thus supporting the reproductive stress hypothesis. Parental body condition during the incubation period was positively related to the probability of successfully fledging young, providing additional support for the reproductive stress hypothesis.     

Repeatability of basal metabolism in breeding female kittiwakes Rissa tridactyla

We studied kittiwakes (Rissa tridactyla) breeding near Ny-Ålesund (79° N, 12° E) on Svalbard. In 1997, the basal metabolic rates (BMRs) of 17 breeding females were measured during the incubation and chick-rearing periods. The mean body mass of the kittiwakes decreased significantly (by 10%) between the incubation and chick-rearing periods. At the same time, both the whole-body and mass-specific BMRs decreased significantly. There was a positive and significant relationship between the BMR residuals from the incubation period and those from the chick-rearing period. Thus, the BMR of incubating female kittiwakes is a significant predictor of their BMR during the chick-rearing period. New BMR data were collected in 1998 from ten of these females, measured around the chick-hatching date. Repeatability values were calculated using either (i) the data for eight individuals for which three BMR measurements existed, or (ii) all the data from both years, yielding significant repeatabilities of 0.52 and 0.35, respectively. These values indicate that between 48 and 65% of the observed variation in BMR is due to intraindividual variability, while between-individual variability accounts for 35 to 52% of the variation in the BMR. This is the first report of a significant repeatability of the BMR of an endothermic organism across an elapsed time of more than one day.     

Components of productivity in black-legged kittiwakes Rissa tridactyla: response to supplemental feeding

In contrast to the high productivity of black-legged kittiwakes in Britain, kittiwakes at many colonies in Alaska have failed chronically to reproduce since the mid 1970s. To determine if food is limiting productivity and, if so, at what stages of nesting food shortages are most severe, in 1996 and 1997 we supplementally fed kittiwakes nesting on an abandoned building. The effects of feeding were stronger in 1997 than in 1996, possibly because naturally occurring prey were of poorer quality in 1997. Consumption of supplemental herring declined as egg laying approached then increased slowly during incubation and more rapidly after hatching. All of the six components of productivity we studied were improved by supplemental feeding to some degree. Supplemental food did not significantly alter laying success in either year, although fed pairs bred at slightly higher rates than unfed pairs in 1997, the poorer food year. In 1996 and 1997, extra food noticeably increased clutch size and hatching success, but significantly so only in 1997. Fledging success and productivity were substantially augmented by feeding in both years. Fed pairs fledged twice as many chicks per nest as did unfed pairs in 1996 and three times as many in 1997. Fed and unfed pairs lost most of their potential productivity through the inability to hatch eggs, and secondarily because of their poor success at raising chicks. The benefits of supplemental feeding did not carry over from one stage of breeding to another. Pairs cut off from supplemental food after laying or hatching performed similarly to pairs that had not been previously fed. This study provides benchmark values of breeding performance attainable by kittiwakes in Alaska under optimal conditions. These values are comparable to highly productive colonies in Britain and suggest that differences in life-history characteristics between Pacific and Atlantic breeding populations are primarily controlled by food supply.     

Evidence for an intrinsic energetic ceiling in free-ranging kittiwakes Rissa tridactyla

1. The rate at which free-living animals can expend energy is limited but the causes of this limitation are not well understood. Theoretically, energy expenditure may be intrinsically limited by physiological properties of the animal constraining its capacity to process energy. Alternatively, the limitation could be set extrinsically by the amount of energy available in the environment or by a fitness trade-off in terms of reduced future survival associated with elevated metabolism.
2. We measured daily energy expenditure (DEE) using the doubly labelled water method in chick-rearing black-legged kittiwakes ( Rissa tridactyla ) at a study site close to the northern limit of their breeding range over 5 years. We measured breeding success, foraging trip duration and diet composition as proxies of resource availability during these years and estimated the probability of parent kittiwakes to return to the colony in relation to their energy expenditure in order to determine whether kittiwakes adjust their DEE in response to variation in prey availability and whether elevated DEE is associated with a decrease in adult survival.
3. We found that DEE was strikingly similar across all five study years. There was no evidence that energy expenditure was limited by resource availability that varied considerably among study years. Furthermore, there was no evidence of a negative effect of DEE on adult return rate, which does not support the hypothesis of a survival cost connected to elevated energy expenditure.
4. The additional lack of variation in DEE with respect to ambient temperature, brood size or between sexes suggests that kittiwakes at a time of peak energy demands may operate close to an intrinsic metabolic ceiling independent of extrinsic factors.     

Natural attachment duration of nymphs of the tick Ixodes uriae (Acari: Ixodidae) on kittiwake Rissa tridactyla nestlings

Daily searches of the plumage of 62 kittiwake (Rissa tridactyla) chicks were carried out for 3 weeks to determine the attachment duration of the tick Ixodes uriae. All ticks recorded were nymphs and their mean duration of attachment was 5.2±1.7 days (n=93). Seventy-four per cent of the ticks survived to engorgement and were assumed to have fed successfully. The mean duration of attachment of successful ticks was 5.8±1.0 days (n=69), significantly longer than that of unsuccessful ticks (3.3±2.1 days, n=24). There was no difference in survival rates to engorgement between ticks attached to feathered and unfeathered parts of the body (78.3 and 79.2%, respectively), but the duration of attachment of successful ticks was significantly longer on unfeathered compared to feathered areas. There were no significant differences in the survival to engorgement or duration of attachment between ticks found on young (c.11 days) and old (>11 days) kittiwake nestlings. The attachment duration was not related to the total number of ticks found on the host or the number of ticks present in the immediate vicinity of the site of attachment. This work provides important data on the parameter of attachment durations of nymphal I. uriae on free-living kittiwakes; the data can be used for incorporation into the application of population modelling.     

Adult survival of Black-legged Kittiwakes Rissa tridactyla in a Pacific colony

Breeding Black-legged Kittiwakes Rissa tridactyla survived at a mean annual rate of 0.926 in four years at a colony in Alaska. Survival rates observed in sexed males (0.930) and females (0.937) did not differ significantly. The rate of return among nonbreeding Kittiwakes (0.839) was lower than that of known breeders, presumably because more nonbreeders moved away from the study plots where they were marked. Individual nonbreeders frequented sites up to 5 km apart on the same island, while a few established breeders moved up to 2.5 km between years. Mate retention in breeding Kittiwakes averaged 69% in three years. Among pairs that split, the cause of changing mates was about equally divided between death (46%) and divorce (54%). Average adult life expectancy was estimated at 13.0 years. Combined with annual productivity averaging 0.17 chick per nest, the observed survival was insufficient for maintaining population size. Rather, an irregular decline observed in the study colony since 1981 is consistent with the model of a closed population with little or no recruitment. Compared to their Atlantic counterparts, Pacific Kittiwakes have low productivity and high survival. The question arises whether differences reflect phenotypic plasticity or genetically determined variation in population parameters.     

云南鸟类新纪录——三趾鸥

2009年2月11日,在云南省昆明市滇池海埂大坝进行鸟类环志工作时,捕获到一只三趾鸥Rissa tridactyla,这是云南省首次发现三趾鸥.该鸟戴上金属脚环后在原地放飞,环号为H04-1583.     

Stage-dependent foraging in breeding black-legged kittiwakes Rissa tridactyla: distinguishing behavioural responses to intrinsic and extrinsic factors

Different phases of the annual cycle in birds and mammals are often associated with characteristic and recurrent foraging behaviours. The extent to which stage‐dependent changes in foraging behaviour are caused by intrinsic or extrinsic factors is unclear. We controlled for the effects of extrinsic factors by synchronising groups of incubating and chick‐rearing black‐legged kittiwakes Rissa tridactyla. Synchrony amongst incubators and rearers was achieved experimentally by switching eggs between nests. Behavioural responses to the treatment varied between the sexes. Male kittiwakes with prolonged‐incubation made fewer foraging trips but of greater duration compared to those rearing chicks resulting in no change in the time spent on trips between the two groups. Females with prolonged‐incubation carried out fewer trips than those rearing chicks but trip duration did not differ between the two stages which resulted in prolonged‐incubating birds spending a lower percentage time on trips. In contrast, foraging ranges did not differ between prolonged‐incubation and chick‐rearing birds for either sex. This suggests that extrinsic factors, such as food availability and distribution determine kittiwake foraging locations and ranges, whereas intrinsic factors, reflected in parental duties, constrain nest attendance. Female prolonged‐incubators invested lower levels of parental effort, in terms of daily energy expenditure, compared to chick‐rearers whereas males did not show stage‐related differences in energy expenditure. This provides evidence that incubation could be an energetically cheaper stage although under normal conditions this difference may be masked by temporal variation in environmental factors. We conclude that while conditions differ between the incubation and chick rearing stages for kittiwakes at this colony, they are not the main factors prompting changes in stage‐related foraging patterns. Intrinsic factors such as sex differences, or behaviours required for each stage of the annual cycle, rather than extrinsic factors related to seasonal environments, are likely to be the main proximate cause of recurring changes in behaviour between breeding stages.     

Population structure in relation to host-plant ecology and Wolbachia infestation in the comma butterfly

Experimental work on Polygonia c‐album, a temperate polyphagous butterfly species, has shown that Swedish, Belgian, Norwegian and Estonian females are generalists with respect to host‐plant preference, whereas females from UK and Spain are specialized on Urticaceae. Female preference is known to have a strong genetic component. We test whether the specialist and generalist populations form respective genetic clusters using data from mitochondrial sequences and 10 microsatellite loci. Results do not support this hypothesis, suggesting that the specialist and generalist traits have evolved more than once independently. Mitochondrial DNA variation suggests a rapid expansion scenario, with a single widespread haplotype occurring in high frequency, whereas microsatellite data indicate strong differentiation of the Moroccan population. Based on a comparison of polymorphism in the mitochondrial data and sequences from a nuclear gene, we show that the diversity in the former is significantly less than that expected under neutral evolution. Furthermore, we found that almost all butterfly samples were infected with a single strain of Wolbachia, a maternally inherited bacterium. We reason that indirect selection on the mitochondrial genome mediated by a recent sweep of Wolbachia infection has depleted variability in the mitochondrial sequences. We also surmise that P. c‐album could have expanded out of a single glacial refugium and colonized Morocco recently.     

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.