Dramatic changes in the return date of Guillemots Uria aalge to colonies in Shetland, 1962–2005

Capsule Changes in return date coincided with marked changes in population size that probably resulted in fluctuating competition for nest-sites.

Aims To document the changes in return dates over a 44-year period and to identify the factors associated with these changes.

Methods We compared changes in return date at Shetland colonies with those for the Isle of May, southeast Scotland, and with the available information on population size, the abundance of some fish species eaten by Common Guillemots and large-scale changes in the oceanography and climate of the eastern Atlantic as reflected by the winter index of the North Atlantic Oscillation (NAO).

Results Common Guillemots normally return to colonies in Shetland in late winter. However, during the 1960s return dates became gradually earlier with birds present from early October. Autumn return remained the norm for about ten years after which return dates gradually reverted back to late winter. In contrast, Common Guillemots on the Isle of May, 400 km south of Shetland, showed no marked shift, returning in October each year. There was a strong negative correlation between date of return of Shetland birds and population size, whereas on the Isle of May birds came back earlier when there was a large positive winter NAO index. There was no convincing evidence that changes in wintering areas or fish abundance influenced when birds returned to the colonies, although the fish data may not have been collected on the correct spatial scale.

Conclusion Competition for high quality nest-sites is the most likely reason for Common Guillemots returning to the colonies during the autumn and winter.     

Site use and fidelity in the Common Guillemot Uria aalge

The site fidelity of 470 colour-ringed Common Guillemots Uria aalge with at least 1 year of breeding experience was followed on the Isle of May from 1982 to 1993. On average, 85.7% of birds recorded breeding at a site in any year were present at the same site in the following season. Of those which did not retain their site, 35% had moved to another site, 25% were nonbreeders and 40% were not seen. Site- fidelity of birds which returned to the colony the next year was 91%. There were no significant age or sex effects, but there were significant (and unexplained) year and area effects. Most birds which changed sites moved less than 2 m. Some birds which obtained a new site improved their nesting success, but many others became nonbreeders; it is speculated that the former chose to move site, whilst the latter had been evicted.     

Survival of adult Common Guillemots Uria aalge at three Scottish colonies

We present the results of a mark–recapture analysis of survival rates of Common Guillemots with at least one year's breeding experience for birds from three Scottish colonies. Estimates of overall survival of adults from the Isle of May (North Sea), Canna and Colonsay (both off northwest Scotland) from an analysis combining data from the three colonies were 94.8% (se = ±0.6), 92.4% (se = ±0.9) and 96.7% (se = ±0.6) respectively. The rates from Canna (but not Colonsay) differed significantly from those of guillemots on the Isle of May. The Canna and Colonsay figures are the first survival estimates for guillemots in northwest Scotland, an area of high conservation concern for this species. Low input, long-term studies of seabirds can produce important data on adult survival.     

Intercolony movement and behaviour of immature Guillemots Uria aalge

A minimum of 61 Guillemots Uria aalge ringed as chicks at other colonies was recorded at the Isle of May between 1987 and 1991 (four prior to 1990, six in 1990, 51 in 1991). Two were adults and 59 were immatures. Most of the British and Irish colonies where many chicks had been ringed were represented. Higher proportions of birds ringed at colonies relatively close to the Isle of May were observed compared to birds ringed at more distant sites. Visitors ranged in age from 2 to 5 years, with single 10- and 14-year-olds. Relatively fewer 2- and 5- than 3- and 4-year-old birds were seen. Visiting birds were usually recorded once only, significantly less often than native immatures of the same age, except for 2-year-olds. The proportion of birds occupying sites on intertidal rocks as against sites in the colony appeared to be higher compared with native birds of the same age. Visiting birds formed a substantial proportion of the immature population. Ten immature birds from the Isle of May were recorded elsewhere. Four of these had been seen on the Isle of May previously, and four were seen there subsequently. A 6-year-old hatched at Sumburgh ( c. 420 km by sea) bred at the Isle of May in 1992.     

A comparison of the biology of the normal and bridled forms of the Guillemot Uria aalge at a single colony

The proportion of Guillemots on the Isle of May showing bridling has remained constant for 49 years. Although clumping of birds occurred, there was no obvious assortment of sexes. We detected no systematic differences in the breeding of bridled and normal individuals.     

Within- and between-year variation in the juvenile survival of Common Guillemots Uria aalge

We studied juvenile survival of 20 cohorts of Common Guillemot Uria aalge chicks colour-ringed on the Isle of May, Scotland, using both live observations at the colony and dead recoveries, allowing estimation of fidelity to the colony as well as survival. In this seabird, chicks leave the colony when only partly grown and are cared for by the male parent for several weeks afterwards. First-year survival varied strongly between cohorts, with a mean of 56% (range 30–91%). We did not identify any covariates which could explain this variation, whether relating to climate, population size or prey density. Survival was low during two regime shift episodes in the North Sea (1987–90 and 2000 onwards). Early hatched chicks were substantially more likely to survive than those hatching later in most years, whereas body condition at ringing had no detectable effect. Ringing recoveries indicated that mortality was highest in mid-winter, i.e. well after the cessation of paternal care. These results do not support the hypothesis that variation in prey quantity or energy content before fledging is a primary driver of variation in juvenile survival. Rather, it seems that chicks of high-quality parents are more likely to survive, as high-quality females tend to lay earlier in the season, and high-quality males presumably are better able to prepare their chicks to survive their first winter at sea. Very few (4%) Guillemots emigrated permanently before age 3 years, but from age 5 onwards 25–30% of birds annually left the colony or otherwise became unobservable.     

Fish fed to young Guillemots, Uria aalge, and used in display on the Isle of May, Scotland

Guillemots on the Isle of May fed their young on Sandeels and Sprats, the proportion of the latter being highest late in the day and the season. There was a peak of feeding after dawn when many large Sandeels were brought. Weather had no effect on feeding rate but this did increase with the chicks age. Food appeared to be abundant. A large Sprat was the most efficient return for effort and Guillemots may select the most energy-rich prey. Display fish (mainly small Sandeels) were brought throughout the season, mostly by females which often ate them later. The significance of the fish-carrying display is obscure.     

Increase in frequency of the bridled morph of the Common Guillemot Uria aalge on the Isle of May, 1946–2000: a return to former levels?

Bridling in the Common Guillemot Uria aalge is a classic example of a stable ratio-cline polymorphism. Between 1946 and 2000 the frequency of the bridled morph of the Common Guillemot colony on the Isle of May, south-east Scotland, increased significantly from 3.5% to 5.9%. Demographic data collected between 1982 and 2000 indicated that the average breeding success of a pair including at least one bridled bird was 83.8%, significantly greater than 79.5% in unbridled pairs. Over the same period the average overwinter survival was 96.3% for bridled birds and 94.7% for unbridled birds, but the difference was not statistically significant. A population model showed that the increase in frequency of bridling could be accounted for as a return to an equilibrium level close to the 5.04% observed in 1936. Differences in breeding success contributed relatively little to the increase, which could be explained without recourse to differential selection pressure. The latitudinal variation in bridling suggests that bridled morphs are more cold tolerant than unbridled ones. However, the increase in the frequency of bridled birds on the Isle of May was not associated with any decrease in sea or air temperatures over the period.     

The Breeding of the Dipper in North Westmorland

In 1983–84, 14 radio transmitters were attached to Guillemots on the Isle of May. Those with internal tuned loop transmitting aerials, glued in the centre of the bird's back with a small amount of resin were most satisfactory. Most birds retained their transmitters for 10–20 days. The bird's behaviour was sometimes altered and ways of minimizing this are discussed. Except when they had young, three (apparently normal) breeding Guillemots spent very little time on the sea in front of their site. The maximum foraging range was smallest at this time.     

The breeding biology of Guillemots Uria aalge on the Isle of May over a six year period

The breeding of Guillemots was studied in five areas of different breeding density and habitat type on the Isle of May in 1981-86. Prior to 1981 numbers were increasing at 5 6° per annum but during the study the rate of increase slowed down and from 1983 to 1986 numbers were fairly constant. Adult survival was high, with a mean minimum annual adult survival of 930% (s.e. = 03). Observations in 1986 suggested that the percentage return of colour-marked immature birds was low, with only l-6% and 5.5% of second and third year birds being seen. We suggest that poor recruitment was responsible for the levelling off in numbers at the colony.
The timing of laying was constant from year to year in 1981-85 but was later in 1986. It was significantly and inversely related to sea temperature the previous March. There was a consistent ranking in median laying dates amongst the areas, with area 1 (the highest density of birds) always earliest. However, there was no significant difference in synchrony between the areas. Overall breeding success was high (0–71-0-82 young fledged per pair). There was no consistent ranking of breeding success with breeding density, habitat type or laying synchrony.
The only aspect of Guillemot biology which changed significantly was the daily food intake of a chick which approximately halved during the study period. However, this reduction in food intake had no detectable effect on either the weight of chicks with wing lengths greater than 60 mm or the amount of time off-duty breeders spent at the site. Both of these parameters were still consistent with conditions being favourable in 1986.     

Increased adult mortality and reduced breeding success with age in a population of common guillemot Uria aalge using marked birds of unknown age

Although there is general consensus about the existence of senescence in vertebrates, empirical evidence of senescence in demographic parameters in wild populations is limited. Data on breeding success and survival of breeding common guillemots Uria aalge were collected over 20 years on the Isle of May (Scotland) using a pool of individuals marked as adults. Because the years of hatching of individuals were not known, we used the time (years) elapsed since first capture (TFC) as a measure of age. The use of this proxy did not create any bias in estimating senescence in the case of a linear decline, nor did it greatly decrease the power of a test for senescence. Breeding success declined significantly from 0.81 (95% CI: 0.77–0.84) to 0.62 (0.54–0.68) over the study period. It also varied in relation to age, initially increasing (from 0.62 (0.54–0.68) at TFC of 0 year to 0.76 (0.73–0.79) at TFC of 9 years) up to a plateau (from TFC of 9 years with 0.76 (0.73–0.79) until TFC of 13 years with 0.77 (0.74–0.79) before declining in later life to 0.70 (0.61–0.78) at TFC of 19 years). Survival was generally high and varied significantly from year to year. It also declined with TFC: survival of birds marked in 1982 decreased from 0.92 (0.85–0.96) at a TFC of 0 year to 0.88 (0.82–0.92) at a TFC of 19 years. Resighting probabilities also declined with TFC suggesting that the oldest birds do not come back to the colony to breed as regularly as younger individuals. These findings indicate that individual common guillemots on the Isle of May showed both actuarial and reproductive senescence.     

To breed or not to breed: causes and implications of non-breeding habit in the willow tit Parus montanus

Causes and consequences of non-breeding in willow tits were studied in northern Finland during 1986–1992. The breeding status was sex and age biased; males and yearling birds were in excess among the non-reproducers. Due to sex bias in the population it appeared detrimental for males to lose a mate, especially shortly before breeding. Lack of a mate was a important factor for males not reproducing (37% of non-breeding males) than for females (14%). Most of the non-breeding birds maintained a pair bond which only rarely broke up for the next breeding season (divorce rate 5.5%). This implies that parental incompatibility is not a possible explanation for pairs not reproducing. Males that did not breed tended to survive better than reproducing ones, whereas such a relationship was not found for females. It is possible that this sex-related difference in survival cost is attributable to quality differences among non-breeding individuals. It was especially low-quality yearling females, with low survival prospects, that were responsible for the discrepancy. The proportion of non-breeding females in the population correlated highly with clutch size and subsequent juvenile survival. It is therefore suggested that for most of these females non-breeding is a phenotypic response to low offspring value in the prevailing circumstances (inter-generational tradeoff). However, it is uncertain whether willow tits in a northern population can use breeding density as an indicator of changing survival prospects of their descendants, as suggested by Ekman and Askenmo (1986) for southern Sweden.     

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.     

Age-related effects on breeding phenology and success of Common Guillemots Uria aalge at a North Sea colony

Capsule: Common Guillemots Uria aalge show delayed breeding and marked age-related changes in reproductive success consistent with improved performance with experience.

Aims: To determine age of first breeding and age-related effects on breeding phenology and success of Common Guillemots.

Methods: Resighting data from a long-term colour-ringing study of Common Guillemot chicks were combined with observations of breeding phenology and success to follow the recruitment process, breeding phenology and success of 62 birds at a major North Sea colony over a 30-year period.

Results: The median age of first breeding of Common Guillemots was 6.6 years. There were no detectable costs of first breeding on return rates or the likelihood of breeding the next season but first time breeders bred later and less successfully. Age of first breeding and lifetime breeding success both varied among individuals but there was no clear optimal age of first breeding and early first breeding was not associated with higher lifetime breeding success.

Conclusions: Common Guillemots in the Isle of May population delayed breeding for 3–4 years beyond physiological maturity. The marked increase in breeding success with age was consistent with improved performance with experience rather than selection for higher quality individuals. Findings from this study will inform population models by providing improved estimates of age of first breeding and age-related changes in reproductive performance.     

Seasonal declines in incubation periods of Brünnich's Guillemots Una lomvia:testing proximate causes

In many species of bird, eggs laid late in the laying period hatch after a shorter incubation period than do eggs laid early. These seasonal declines in incubation period are generally thought to confer evolutionary advantages, but the proximate mechanisms that underlie them are poorly understood. Seasonal declines in incubation period are usually attributed to: (1) seasonal increases in ambient air temperatures; (2) seasonal changes in the behaviour of incubating birds; and/or (3) seasonal declines in egg size. In a previous study, Common Guillemot Una aalge incubation periods declined with laying date at a low-Arctic colony. As there was no support for hypotheses 1 or 2, it was suggested that this occurred because egg size declined with laying date, but eggs were not measured in that study. We recorded similar seasonal declines in the incubation periods of the single eggs laid by Brünnich's Guillemots Una lomvia at two low-Arctic colonies in four years. Neither seasonal variation in ambient air temperatures, nor in the behaviour of incubating adults, appeared to cause the declines. As predicted for Common Guillemots, incubation period increased with egg size among Briinnich's Guillemots, in one of two years. However, incubation period declined with laying date in the absence of corresponding declines in egg size. We conclude that none of the three commonly proposed proximate mechanisms adequately explains the seasonal variation in guillemot incubation periods. Several testable, alternative mechanisms are explored.     

The population dynamics of the Eider Duck Somateria mollissima and evidence of extensive non-breeding by adult ducks

The dynamics of an Eider Duck population have been investigated over 25 years, using census and capture-mark-recapture methods. During the study the population increased two and a half fold, with two periods of major increases in numbers, giving a stepped growth pattern. Mean clutch size showed considerable annual variation, the extremes being 5.40 and 3.78 eggs. The variation in clutch size was greater than that recorded in the Netherlands. Adult female Eiders had a high annual survival rate, averaging 0.895, and varying between 0.75 and 1.00 in individual years. The survival rate decreased markedly in the old ducks. There was no indication of any change in the survival rate during the study. Recruitment of ducks to the breeding group was irregular, with most years showing little recruitment and a few years high recruitment. However, recruitment, associated with good duckling survival, appears to have been the main factor associated with increase in the population. In many years, an appreciable proportion of the surviving ducks, which had already bred in a previous year, failed to nest. The extent of non-breeding increased during the study and in one year, 1973, this reached 65%. Lower clutch size and adult survival were associated with years of high non-breeding. The ‘red-tides’ in 1968 and 1975 appeared to have little effect on the Eider. It is suggested that the Eider missed breeding in years in which its survival was potentially poor, in order to maximize its reproductive output during its life span. This is supported by the smaller clutch size laid by those females which nest in years when many females fail to breed. It is suggested that young ducks may also miss breeding in the year after first nesting but this is not associated with the non-breeding in older ducks, although it may he related to body condition. It is suggested that non-breeding by adults of long-lived birds may be widespread. This has important implications in survey work based on nest counts.     

Covariation among demographic and climate parameters in whiskered auklets Aethia pygmaea

Annual survival rate and other demographic parameters of whiskered auklets Aethia pygmaea , a small planktivorous seabird, were measured at Buldir Island Alaska during 1992–2003 to provide comparative information for auk life history studies and to test for links among climate, age, productivity and survival. Using two 9  m mistnets, we captured and recaptured 384 adult and 193 sub-adult (one year old, a known-age component of our sample) birds as they arrived at the colony after dark during May and June (1,730  capture events). The best fitting model indicated a lower initial survival rate over the first year following marking (0.708±0.036 SE), and subsequent survival (mean 0.835±0.029) covarying with the Aleutian Low Pressure climate Index, with higher auklet survival in years with weak low pressure over the Aleutian Islands. Annual survival rate varied from 0.726±0.127 in 1998–99 to 0.994±0.077 in 1994–95, rates similar to those previously reported for least A. pusilla and crested auklets A. cristatella . A model based only on recaptures of known-age birds indicated a lower local survival estimate over the first year following marking (age one to two years), with no other age-effects on survival. Breeding propensity by age inferred from recaptures of birds with fully-developed brood patches that were originally marked as sub-adults (one year olds) indicated 53% breeding at age two, 94% breeding at age 3, 97% breeding at age 4 and 100% breeding thereafter. The sex ratio of the sampled birds was significantly male biased (60/40), likely due to differences in behaviour between males and females during the incubation stage. Taken together, our data indicate that whiskered auklet survival and productivity covaried with continuous variation in large-scale climatic conditions, the mechanism being either negative effects of stormy North Pacific weather or indirect effects on food supplies.     

Recruitment to a seabird population depends on environmental factors and on population size

1. A novel capture-mark-recapture (CMR) method was used to build a multistate model of recruitment by young birds to a breeding population of common guillemots Uria aalge on the Isle of May, Scotland. Recruitment of a total of 2757 individually marked guillemots over 17 years was modelled as a process where individuals had to move from an unobservable state at sea, through a nonbreeding state present in the colony, to the breeding state. The probabilities of individuals returning to the colony in a given year, at age 2 and 3-4 years, were positively correlated with an environmental covariate, the winter North Atlantic Oscillation index (WNAO) in the previous years. 2. For 2 year olds, there was a negative relationship with breeding population size, suggesting that density dependence operated in this colony through limitation of food or some other resource. 3. Survival over the first 2 years of life varied with cohort, but was unrelated to the WNAO. Mean survival over this 2-year period was high at 0.576 (95% CI: 0.444; 0.708). 4. This high survival, combined with a low 'local' survival after age 5 years of 0.695 (0-654; 0.733) and observations of Isle of May chicks at other colonies, suggests that most surviving chicks return to the natal colony before deciding whether to recruit there or move elsewhere.     

Effects of prey abundance on the foraging behaviour, diving efficiency and time allocation of breeding Guillemots Uria aalge

The foraging behaviour of Guillemots Uria aalge at sea was compared between 2 years of radically different food abundance. Radio telemetry was used to determine foraging locations and diving patterns. In the poor compared with the good food year, foraging trips were much longer, the birds foraged more than six times further from their breeding sites, they spent over five times as much time diving when at sea and their estimated energy expenditure was twice as great. Time spent foraging in the poor food year was at the expense of time spent sitting at the colony. The duration of a foraging trip was a poor indicator of distance travelled but a good indicator of the amount of time spent diving. Mean dive durations, surface pause durations and interbout periods did not differ between years, but individuals made more than four times as many dives per diving bout in the poor food year. Surface pause lengths did not vary with water depth in either year. In the poor food year, birds made shorter surface pauses for a dive of a gi^ven duration than in the good food year, possibly accepting a lactic acid debt in order to maximize searching time, The duration of the interbout period was positively related to the number of dives in the previous bout, and dives tended to get shorter in long diving sequences, suggesting possible exhaustion effects. These data demonstrate that breeding Guillemots have the capacity to adjust their foraging behaviour and time budgets in response to changes in food abundance, but this flexibility was not sufficient to compensate fully for the very low food abundance experienced by birds in this study.     

An Experimental study of reduced parental effort and future reproductive success in the puffin, Fratercula arctica

1. The puffin, a long-lived seabird, was studied on the Isle of May, East Scotland between 1990 and 1992. During two of these years, parental effort was experimentally decreased by supplementary feeding of young. This aimed to identify inter-year reproductive costs, and show whether they took the form of reduced adult survival, reduced fledging success and/or a reduction in the 'quality' of offspring in the following year.
2. The feeding treatment significantly reduced the daily number of feeds delivered by experimental parents by 67% in 1990 and 87% in 1991.
3. The proportions of experimental and control parents returning to the colony in the year following manipulation did not differ significantly, although in 1991, 2·5 times as many controls (young unfed) as experimental birds (young fed) failed to return.
4. The fledging success of experimental pairs in the year following manipulation (68%) was significantly higher than that of controls (24%).
5. Experimental pairs raised young with significantly higher body condition (Residual Peak Mass) than that of controls in the year following manipulation (1992).
6. Experimental parents did not differ from controls in their body condition (Lipid Reserve Mass) or rate of reserve depletion, either in the year of manipulation or in the following breeding season; hence there was no evidence for a role of the measured component of body condition in the cost mechanism.
7. The study demonstrated inter-year reproductive costs for puffins and supported the hypothesis that long-lived species reduce the 'quality' of their offspring or abandon a breeding attempt rather than compromise their survival and future opportunities to reproduce.