Population status of Common Guillemots Uria aalge at a colony in western Alaska: results and simulations

Colony-wide boat-based censuses suggest that numbers of Common Guillemots at Bluff, Alaska declined markedly in the late 1970s and remained low until 1983. Annual land-based counts of study plots since 1979 generally showed no trend. However, numbers declined substantially in one study area between 1978 and 1983. Reproductive success varied considerably among years and was highest in moderately warm summers. Simulation analyses suggest that the changes in numbers cannot be attributed to changes in reproductive success several years previously. Thus, factors affecting survival, presumably in winter, are apparently implicated in the declines.     

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.     

The effects of food abundance on breeding performance and adult time budgets of Guillemots Uria aalge

The breeding performance, food fed to chicks and adult time budgets of Guillemots Uria aalge were examined in a year of high and a year of low food availabiIity. There was no difference between the 2 years in reproductive success, although the rate of chick feeding, chick weight and fledging success were greater in the year of high food availability. On average, chick prey items were larger in the poor food year, but this was insufficient to compensate for the lower feeding frequency. Chick feeding frequency did not differ between days in the good year but did increase later in the season in the poor food year. Compared with the high food availability year, adult Guillemots in the year of low food availability spent much less time resting at the breeding colony. and their foraging trips were twice as long. Foraging birds tended to make several successive trips before resuming brooding duties from their mates when food supplies were good, but in the low food availability year single trips were the norm. These results demonstrate that predators experiencing reduced food supply may mitigate the effects on their reproductive output by shifting their time allocation such that more time is available for foraging.     

Among‐colony synchrony in the survival of Common Guillemots Uria aalge reflects shared wintering areas

Spatiotemporal variation in survival may be an important driver of multi‐population dynamics in many wild animal species, yet few scientific studies have addressed this issue, primarily due to a lack of sufficiently comprehensive and detailed datasets. Synchrony in survival rates among different, often distant, subpopulations appears to be common, caused by spatially correlated environmental conditions or by movement of animals from different sites such that their ranges overlap. Many seabird populations are effectively isolated during the breeding season because colonies are widely separated, but over the winter, birds disperse widely and there may be much mixing between different populations. The non‐breeding season is also the period of main mortality for seabirds. Using mark–recapture and ring‐recovery data, we tested for spatial, temporal and age‐related correlations in survival of Common Guillemots Uria aalge among three widely separated Scottish colonies that have varying overlap in their overwintering distributions. Survival was highly correlated over time for colonies/age‐classes sharing wintering areas and, except in 2004, was essentially uncorrelated for those with separate wintering areas. These results strongly suggest that one or more aspects of the winter environment are responsible for spatiotemporal variation in survival of British Guillemots, and provide insight into the factors driving large‐scale population dynamics of the species.     

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.     

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.     

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.     

Geolocators reveal an unsuspected moulting area for Isle of May Common Guillemots Uria aalge

Capsule Data from geolocators deployed on adult Common Guillemots from a colony in southeast Scotland indicated that they normally winter in the North Sea up to 1000?km southeast of the colony. However, one bird unexpectedly moved 3000?km northeast to moult in the Barents Sea.     

The importance of observer effort on the accuracy of breeding success estimates in the Common Guillemot Uria aalge

ABSTRACT

Capsule: A data-thinning approach was used to assess the effects of reducing the frequency of nest-checks on estimates of breeding success of Common Guillemots Uria aalge. Inter-year and inter-colony differences in fledging age and their implications for setting a minimum age after which a chick could be assumed to have fledged were evaluated.

Aims: To assess the consequences of reducing sampling frequency on the estimation of breeding success, and on the robustness of the assumption that breeding has been successful if a chick survives to 15 days old.

Methods: Breeding success, ages at fledging and loss of chicks were estimated from daily checks at two Scottish colonies over a six-year period. Data-thinning was used to assess the consequences of reducing checks from daily to every two or three days. Breeding success was recalculated assuming that all chicks surviving to 15 days fledged.

Results: Reducing the frequency of checks from daily to every second or third day resulted in a small but statistically significant increase in the estimate of success. Between 20% and 25% of chick losses occurred when chicks were ≥15 days old. Assuming that these chicks had fledged resulted in significant increases in breeding success.

Conclusion: Assumptions about whether or not a chick fledged had a greater impact on the estimate of breeding success than reducing the frequency of nest-checks from daily to every second or third day. There was no threshold age after which a chick could be assumed to have fledged. These findings are relevant to other monitoring schemes where there is pressure to reduce input. Sampling methods used in monitoring schemes need to be clearly stated and changes in protocols documented so that sampling effects can be incorporated into future analyses.     

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.     

Variation in the correction factor used for converting counts of individual Guillemots Uria aalge into breeding pairs

The correction factor k (breeding pairs/total count of birds) was measured for two nesting groups of Guillemots Uria aalge over 6–7 years. The value varied between different subgroups but there were no significant differences between groups or between years. These results, and those from other studies, suggest that a value of 0–67 is applicable to a range of colonies. Values of k for Razorbills Aha tarda were more variable. A plea is made for the presentation of original counts rather than 'corrected' figures in published work.     

Survival and non-breeding of adult Common Guillemots Una aalge

Annual survival of colour-ringed Common Guillemots Uria aalge with at least 1 year of breeding experience was measured in five study areas on the Isle of May, southeast Scotland from 1982 to 1993. Annual survival averaged 94.9%. There were no significant sex, year or area differences. Between 5% and 10% of the birds known to be alive in any year did not breed, and on average such birds appeared to survive less well (87%) than breeders (95%). Non-breeding did not occur randomly within the population, and 6% of birds accounted for 47% of the cases of non-breeding. It appeared that non-breeding in Common Guillemots on the Isle of May was socially induced with most cases occurring after an individual had lost its site.     

Nest site characteristics, duration of use and breeding success in the Guillemot Uria aalge

The fate of all eggs laid by Guillemots Uria aalge in seven study areas on the Isle of May was recorded in 12 consecutive seasons. The average success at each site was related to its physical characteristics, bird density, position on the cliff and availability of suitable habitat for ticks. The numbers of neighbours and walls, type of site, slope where the egg was incubated and distance from the top of the cliff all had a significant effect on both hatching and breeding success. Potential tick habitat had no significant effect. Much variation in hatching and breeding success remained unexplained. Sites used when the colony was less than half its current size and most frequently in the period 1984–1995 were the most successful. Since an average Guillemot pair remains together at a site for only 3–4 years. the consistently high success of many sites over a period of 10–12 years suggests that the most used sites were occupied by a succession of high-quality birds.     

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.     

The visual fields of Common Guillemots Uria aalge and Atlantic Puffins Fratercula arctica: foraging,vigilance and collision vulnerability

Significant differences in avian visual fields are found between closely related species that differ in their foraging technique. We report marked differences in the visual fields of two auk species. In air, Common Guillemots Uria aalge have relatively narrow binocular fields typical of those found in non‐passerine predatory birds. Atlantic Puffins Fratercula arctica have much broader binocular fields similar to those that have hitherto been recorded in passerines and in a penguin. In water, visual fields narrow considerably and binocularity in the direction of the bill is probably abolished in both auk species. Although perceptual challenges associated with foraging are similar in both species during the breeding season, when they are piscivorous, Puffins (but not Guillemots) face more exacting perceptual challenges when foraging at other times, when they take a high proportion of small invertebrate prey. Capturing this prey probably requires more accurate, visually guided bill placement and we argue that this is met by the Puffin's broader binocular field, which is retained upon immersion; its upward orientation may enable prey to be seen in silhouette. These visual field configurations have potentially important consequences that render these birds vulnerable to collision with human artefacts underwater, but not in air. They also have consequences for vigilance behaviour.     

The post-fledging survival of young Guillemots Uria aalge in relation to hatching date and growth

The post-fledging survival of a total of 1277 young Guillemots Uria aalge ringed in 6 years was assessed using sightings of 267 individuals back at the natal colony and recoveries of 46 ringed birds. In two years there was a significant decline in survival prospects with estimated hatching date, the first time such a trend has been demonstrated in the Alcidae. In these years pairs breeding early had a 2–3 times greater chance of having a young survive to return to the colony than pairs breeding three weeks later. Hatching date had no effect on survival prospects in the other four seasons and in none of the years did chick body condition have a demonstrable effect on post-fledging survival.     

Distribution of the bridled form of the Common guillemot Uria aalge in the North Atlantic

This study documents the fourth survey (198 1–1982) of bridled guillemots and provides the most complete information yet available on the distribution of the bridled form of the Common guillemot. The distribution has remained largely unchanged since the last survey (1959-60), although marked changes have occurred at a few colonies. Sea surface temperature "explains" only 35% of the variance in the proportion of bridled guillemots. Minimum air temperatures during the breeding season are much more closely correlated with bridling and "explain" 60% of the variance. No relationship exists between sea temperatures and the proportion of bridled guillemots in wintering areas. The distribution of bridled guillemots may represent the slow, southward spread of the bridled form. Alternatively, selection may favour the bridled form in cooler areas. There is insufficient evidence to distinguish between these two hypotheses, but if selection is important, the correlations with environmental factors suggests that it operates during the breeding season rather than at other times.     

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.