Parallel declines in survival of adult Northern Fulmars Fulmarus glacialis at colonies in Scotland and Ireland

Assessing broad‐scale changes in seabird populations across the North Atlantic requires an integration of available datasets to understand the spatial extent of potential drivers and demographic change. Here, we compared survival of Northern Fulmars Fulmarus glacialis from a Scottish and an Irish colony from 1974 to 2009. Despite lower recapture probabilities of monel‐ringed Irish birds compared with colour‐ringed Scottish birds, survival probability decreased at both colonies. The extent to which the decline in survival is related to density‐dependent processes or other external drivers remains uncertain, but our results suggest that these changes in survival are possibly indicative of larger‐scale processes and are not confined to local colony dynamics.     

Breeding status, contaminant burden and helminth parasites of Northern Fulmars Fulmarus glacialis from the Canadian high Arctic

We studied the relationship between contaminant concentrations in livers, endohelminth prevalence and physiological indices of chronic stress (spleen size, heterophil/lymphocyte ratios) in Northern Fulmars Fulmarus glacialis collected during the breeding season in Nunavut, Canada. No blood parasites were found, similar to reports for other petrel species elsewhere. However, 54% of Fulmars had gastrointestinal helminths, principally cestodes (52% prevalence, mean intensity of 11 worms), nematodes (34% prevalence, 3.6 worms) and acanthocephalans (3%, eight worms). Both prevalence and intensity of helminth infections were lower for Arctic Fulmars than for Fulmars and other petrels found in southern parts of the species' range. Spleen size was not significantly related to either contaminant concentration or presence of parasites, suggesting that Fulmar health was generally unaffected by contaminant and parasite levels at the colony. However, we cannot rule out the possibility that Fulmars with higher parasitaemias or contaminant loads were in poorer condition and did not attend the breeding colony.     

Provisioning and growth rates of nestling Fulmars Fulmarus glacialis: stochastic variation or regulation?

The pattern of chick feeding in the Fulmar Fulmarus glacialis at St Kilda, Scotland, was examined by repeated weighing of chicks throughout 14 consecutive days during the first half of the chick-rearing period in 1994. After correcting for metabolic weight losses, the sizes of positive mass increments between weighings were used to assess meal sizes and feeding frequency for each chick. Individual meals fed to chicks averaged 80.8 g (s.d. ± 21.0 g), or approximately 10% of adult mass. Each chick received 0 to 4 meals per day, with an average of 1.9 meals per chick per day, giving an average interval of around 25 h between meals delivered by each parent. The distribution of time intervals between feeds for each chick (whether single or double meals) followed a negative exponential function with a maximum value of 80 h. These results are not compatible with the idea that the purpose of large fat deposits in procellariiform chicks is to guarantee survival over long intervals between feeds. Over 14 days, the chicks' mean daily food requirements for zero-growth increased from 98 g to 160 g. This corresponded with an increase in feeding frequency but not meal size. Chicks with lower scores for body condition after feeding by both parents received more meals during the subsequent 16 h and had shorter intervals to the next feed, indicating that adults regulated feeding frequency in accordance with chick condition at the previous feed. This does not agree with the hypothesis that lipid accumulation by nestling Procellariiformes is a response to stochastic variation in food delivery associated with an absence of regulation. In view of the diversity of growth and feeding patterns present among the Procellariiformes, it is possible that lipid accumulation in this group does not have a unitary explanation.     

Diets and feeding of Fulmars Fulmarus glacialis during the breeding season:a comparison between St Kilda and Shetland colonies

Diets and nest attendance of Fulmars at Foula, Shetland and St Kilda, Outer Hebrides were examined during the breeding season. At Foula, sandeels formed the bulk of the diet, with fish offal and pelagic zooplankton of minor importance. At St Kilda, 71% of regurgitates consisted of pelagic zooplankton, probably captured at night. Dietary overlap between the two colonies was 14% by species composition. Foraging trips from Foula shortly after chick hatching generally lasted for less than 10 h, while trips from St Kilda late in chick rearing often lasted more than 24 h. The diurnal pattern of feeding suggested mainly nocturnal foraging from St Kilda but both diurnal and nocturnal foraging from Foula.     

Delimiting shades of gray: phylogeography of the Northern Fulmar,Fulmarus glacialis

The Northern Fulmar (Fulmarus glacialis) is a common tube‐nosed seabird with a disjunct Holarctic range. Taxonomic divisions within the Northern Fulmar have historically been muddled by geographical variation notably including highly polymorphic plumage. Recent molecular analyses (i.e., DNA barcoding) have suggested that genetic divergence between Atlantic and Pacific populations could be on par with those typically observed between species. We employ a multigene phylogenetic analysis to better explore the level of genetic divergence between these populations and to test an old hypothesis on the origin of the modern distribution of color morphs across their range. Additionally, we test whether mutations in the melanocortin‐1 receptor gene (MC1R) are associated with dark plumage in the Northern Fulmar. We confirmed that mitochondrial lineages in the Atlantic and Pacific populations are highly divergent, but nuclear markers revealed incomplete lineage sorting. Genetic divergence between these populations is consistent with that observed between many species of Procellariiformes and we recommend elevating these two forms to separate species. We also find that MC1R variation is not associated with color morph but rather is better explained by geographical divergence.     

Breeding status influences timing but not duration of moult in the Northern Fulmar Fulmarus glacialis

Seabirds are key marine top predator species that are often used as indicators of the environmental quality of the oceans. Their breeding phenology has been studied extensively, but their pelagic habits mean less is known about the phenology of other events during the non-breeding period. Here, we used miniaturized saltwater immersion light-based geolocators (GLS) to investigate moult phenology in individuals with known breeding histories in a population of Northern Fulmar Fulmarus glacialis in Orkney, Scotland. As seabirds spend more time on the water during moult, moulting periods can be identified from patterns of variation in the amount of time that birds are in contact with saltwater. Estimates of daily variation in this behaviour during the non-breeding period were based upon wet/dry sensors and then modelled to characterize the timing of the moult. Light-based geolocation provided information on the areas used by each individual during its moult period. Inter-individual variability in moult timing was investigated in relation to sex and breeding success in the previous summer. We found a sex difference in the location of the moult, but not in its timing. However, the timing of moult did differ between individuals that had succeeded or failed in their previous breeding attempt, with successful breeders moulting the latest. In contrast, the duration of moult did not depend on prior reproductive success, but there was evidence of inter-annual variation in moult duration. GLS studies have provided a step change in our understanding of the at-sea distribution of pelagic seabirds. Our work highlights how activity data from these devices can add value to such studies by identifying key phases of the annual cycle, and locations at these times, when seabirds may be at particular risk. Furthermore, our findings indicate that individual and inter-annual variation in breeding success may influence phenological patterns in other phases of the Northern Fulmar annual cycle.     

Weight loss of Antarctic Fulmars Fulmarus glacialoides during incubation and chick brooding

Weight losses incurred by incubating birds of both sexes, and weight changes at the beginning and end of shifts between successive incubation and brooding shifts were studied in the Antarctic Fulmar Fulmarus glacialoides in Adélie Land, Antarctica. The overall rate of weight loss during incubation fasts was 4.46% per day in both sexes but 3.91% per day after the first 2 days of fast once the stomach contents had been digested. Antarctic Fulmars appear to have rates of body-weight loss significantly greater than the rates of weight loss observed in any other species of albatross or petrel. The validity of methods using the rate of weight loss to estimate incubation metabolic rates is discussed, and it appears that for birds incubating for periods of less than 2–3 days the digestion of the stomach contents can bias the estimation. The absence of change in weight at the beginning of shifts during incubation and brooding, together with the short time spent on foraging trips, suggests that food availability is high and/or food is readily accessible to Antarctic Fulmars. The availability of food allows the Antarctic Fulmar to compensate for the high energy cost of breeding that could be due to the cold Antarctic environment and perhaps also to a higher intrinsic metabolism.     

Postnatal development of northern fulmar chicks, Fulmarus glacialis

The slow growth and large fat stores characteristic of many pelagic seabird chicks were generally assumed to reflect infrequent and unpredictable food provisioning by parents. Much less attention has been focused on the importance of intrinsic physiological processes in shaping patterns of development. In this study, we examined postnatal growth and changes in water content of different organs in fulmar chicks, Fulmarus glacialis, from Fair Isle, United Kingdom. After correcting for body size, mass growth rate was as high as in inshore-feeding species, which did not support the notion of an external constraint on growth imposed by the unpredictability of pelagic prey. Pectoral muscles and plumage grew more rapidly than other tissues. Pectorals also had a high water index, probably indicating slower maturation compared with leg muscles, which need to generate heat earlier on to free adults from brooding requirements. Lean dry mass of liver, kidney, and gut decreased markedly toward fledging, presumably because of high energetic costs of maintaining large metabolic machinery in older chicks and analogous to the situation in adult waders before migration. These results suggest that the general pattern of development of fulmars may be linked to changes in resource allocation as chicks grow and possibly a compromise at the tissue level between cell division and the attainment of mature function.     

Breeding strategies of Antarctic Petrels Thalassoica antarctica and Southern Fulmars Fulmarus glacialoides in the high Antarctic and implications for reproductive success

Breeding strategies of two closely related fulmarine petrels were studied on Ardery Island, on the continental coast of East Antarctica, where short summers are expected to narrow the time-window for reproduction. Both species had a similar breeding period (97 days from laying to fledging) but Antarctic Petrels Thalassoica antarctica bred up to 16 days earlier than Southern Fulmars. During the pre-laying exodus, all Antarctic Petrels deserted the colony, whereas some Southern Fulmars Fulmarus glacialoides remained. Antarctic Petrels exhibited stronger synchronization in breeding, made longer foraging trips and spent less time guarding their chicks than Southern Fulmars. Overall breeding success of both species was similar but failures of Antarctic Petrels were concentrated in the early egg-phase and after hatching, when parents ceased guarding. Southern Fulmars lost eggs and chicks later in the breeding cycle and so wasted more parental investment in failed breeding attempts. Different breeding strategies may be imposed by flight characteristics; Southern Fulmars are less capable of crossing large expanses of pack ice and need to delay breeding until the sea ice retreats and breaks up. However, due to the short summer they risk chick failure when weather conditions deteriorate late in the season.     

Das Meeressegeln des Eissturmvogels(Fulmarus glacialis)

Zusammenfassung Der Eissturmvogel zeigt die typische Flugweise des Albatros, die vielfach als dynamischer Segelflug bezeichnet wird. Unsere Beobachtungen über die Einzelkomponenten des Fluges widerlegen jedoch, daß ein ausreichender dynamischer Energiegewinn vorliegt. Es wird angenommen, daß die Vögel die für die Flugmanöver notwendige Auftriebsenergie im niedrigen Teil der Flugbahn gewinnen.

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Summary The Fulmar exhibits the typical manner of flight of the Albatross which has often been designated as dynamic soaring. Our observations on the single components of this flight refute the supposition that there is sufficient yield of dynamic energy. It is supposed that the lift which enables the the birds to make these maneuvers is obtained as the lowest part of the flight.

     

Incubation rhythm in the Fulmar Fulmarus glacialis: annual variation and sex roles

I monitored the incubation schedules of Fulmars Fulmarus glacialis in an Alaskan colony by observing nests where the male and female were of different colour phases. Complete shifts of up to 16 days were recorded; the average shift in mid-incubation was 4–6 days. Mean shift length was inversely correlated with hatching success in 5 years, suggesting that Fulmars adjusted their foraging patterns to annual differences in food availability. Males assumed the larger share (55%) of incubation on average, and a larger share in years with lower hatching success. Serial correlation in the length of incubation shifts had two components—the influence of prior shift lengths on time spent foraging and individual variation. Failure of the male to relieve the female soon after laying resulted in a few breeding failures and egg losses were associated with exceptionally long shifts throughout incubation.     

Static and dynamic expression of life history traits in the northern fulmar Fulmarus glacialis

Understanding the static and dynamic expression of life history traits is a prerequisite for the development of a causal theory of the evolution of aging and of life histories. We analyzed the statics and dynamics of reproduction and survival in a wild population of the northern fulmar, Fulmarus glacialis (Procellaridae). Survival rate is most influenced by year as compared to age and cohort. When temporal variation is ignored, survival rate increases slowly with age and then declines more rapidly at late ages. Survival rate contingent upon reproductive ‘stratum’ (producing an egg, hatching an egg, fledging a hatchling) also exhibits this pattern. Survival and reproduction have a positive static association in that survival rate increases as the apparent energy allocated to reproduction increases (as indexed by stratum). There is a broad distribution of realized lifetime reproductive success, which could be due to ‘fixed’ heterogeneity, with some individuals always having low survival and reproduction and others always having high survival and reproduction, or be due to ‘dynamic’ heterogeneity, with all individuals having the same expected reproductive and survival rates. Analysis of stochastic stratum dynamics indicates that individuals do not remain long in any given stratum and suggest that the variation among individuals with respect to lifetime reproductive success is due to dynamic heterogeneity. The probability of producing an egg increases with age for both sexes, whereas the probability of producing a fledgling initially declines with age and then increases. These results underscore the necessity of understanding the static and dynamic expression of demographic traits when making a causal claim about their evolution.     

Ectoparasites of northern fulmars Fulmarus glacialis (Procellariiformes: Procellariidae) from the Canadian Arctic

We studied the prevalence and intensity of infestation of ectoparasites on northern fulmars (Fulmarus glacialis L.) from a breeding colony in Arctic Canada in June–August 2003. No fleas or ticks were found on any fulmars, but three species of chewing lice (Phthiraptera) were recorded: Ischnocera: Perineus nigrolimbatus (Giebel 1874), Ischnocera: Saemundssonia occidentalis (Kellogg 1896), and Amblycera: Ancistrona vagelli (Fabricius 1787). Non-breeding birds had a higher prevalence of lice than breeding birds, and prevalence varied markedly among louse species. Our study is an important baseline for the occurrence of ectoparasites on northern fulmars in the high Arctic, a region undergoing extensive environmental change due to global warming, and an area where parasites are expected to extend ranges or increase in prevalence under changing annual temperature regimes.     

Competition between two closely related Tetrabothrius cestodes of the fulmar (Fulmarus glacialis L.).

The two congeneic Tetrabothrius cestodes of the fulmar are so closely related that they can only be reliably distinguished by differences in the number of testes. The two species show a considerable distributional overlap along the intestine. Statistically it can be shown that high numbers of the common, anteriorly situated species (T. minor) affect the distribution of the rarer, more posteriorly situated species (T. procerus), causing the latter to occupy a more posterior site in the intestine. Several possibilities are discussed concerning the mechanism by which T. procerus is affected, and how this mildly competitive situation is maintained. It is probable that niche segregation between the two species is more or less complete, but that this is reinforced in concurrent infections by interactive site segregation.     

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.     

Incubation shifts of northern fulmars Fulmarus glacialis in the Canadian high Arctic determined by digital photography

Determining the incubation pattern of northern fulmars (Fulmarus glacialis) at Canadian Arctic colonies is difficult, in part due to challenges with accessing nests and due, in part, to similarities in colour morphs which can be hard to distinguish. We used digital photographs of incubating birds taken ~daily at 49 breeding sites during the second half of the incubation period and compared inter-observer variation in distinguishing incubation shifts by each member of the pair. Using this method gave similar results when different subsets of sites were included and the assessment of incubation shifts by three different people produced similar results, yielding a mean shift duration of 6.4 days. Provided pairs with suitable variation in plumage or bill colour are selected, our results demonstrate that a remote camera technique can provide precise data to assess fulmar responses to annual environmental conditions for feeding and breeding.