Assimilation efficiency of adult Kittiwakes and Brünnich's Guillemots fed Capelin and Arctic Cod

The mean assimilation efficiencies of 10 adult Kittiwakes (Rissa tridactyla) and 10 Brünnich's Guillemots (Uria lomvia) fed on Capelin (Mallotus villosus) were 77.5% and 74.4%, respectively. When fed on Arctic Cod (Boreogadus saida) they were 83.1% and 78.2%, respectively. After correction for nitrogen retention, the assimilation efficiencies decreased to 72.2%, 70.6%, 81.2% and 74.7%, respectively. Kittiwakes and Brünnich's Guillemots seem to have the same ability to utilize the energy of the different food items. The differences in assimilation efficiencies when fed two fish species was mainly related to the fat content of the fish.     

Size,age and diet of polar cod,Boreogadus saida (Lepechin 1773), in ice covered waters

Summary Polar cod (Boreogadus saida) associated with drifting sea-ice were collected in the western Barents sea and north of Svalbard with dip-nets while SCUBA-diving in 1986 and 1987. Length-frequency measurements and otolith-readings suggested that the specimens were either one or two years old. The diet of fish from the western Barents sea (first-year ice) consisted mainly of copepods (Calanus finmarchicus, Calanus glacialis) and the hyperiid amphipod Parathemisto libellula. Fish collected north of the Svalbard archipelago (multi-year ice) had a more diverse diet, in which P. libellula and the sympagic amphipod Apherusa glacialis contributed more to the total diet biomass than copepods.     

Winter diets of four seabird species in the Barents Sea after a crash in the capelin stock

Summary I describe the winter diets (early March) of four seabird species, black-legged kittiwake (Rissa tridactyla), northern fulmar (Fulmarus glacialis), glaucous gull (Larus hyperboreus) and Brüinnich's guillemot (Uria lomvia) collected in the south eastern Barents Sea, in open water approximately 70 km from the ice edge. All species preyed heavily on commercial fish species such as cod (Gadus morrhua), polar cod (Boreogadus saida) and redfish (Sebastes marinus/S. mentella). The median total length of fish eaten ranged from 51 mm in fulmars to 88 mm in Brünnich's guillemots and differed significantly among all species pairs except kittiwake and glaucous gull. Although the size of fish eaten differed among the four bird species they all fed upon much the same age categories of fish. Cod and redfish eaten were almost exclusively one year old, whereas polar cod was 1–4 years old. Other important food items were crustaceans (B unnich's guillemots), and squid Gonatus sp. (fulmars); the glaucous gull also preyed on other birds. Body weights and amounts of stored fat suggest that individuals of all species except Brüinnich's guillemots were in good physical condition. The diets described here differed substantially from those of birds caught mainly during the prelaying season near the colonies in this area. Some of these differences may be attributed to the very low density of capelin (Mallotus villosus) when this study was conducted.     

The occurrence of <Emphasis Type="Italic">Eualus gaimardii gibba</Emphasis> Krøyer 1841 (Crustacea,Decapoda) in the sympagic habitat: an example of bentho-sympagic coupling

We report the first records of a decapod species (Eualus gaimardii gibba) in association with Arctic multiyear sea-ice. Seven specimens inhabiting brine channels in the ice were collected during two field campaigns (September 2003, March/April 2004) by diver operated suction samplers. Salinity tolerance experiments (14–50 PSU) suggest that the species is stenohaline. Gut examinations revealed remains of the sympagic amphipod Gammarus wilkitzkii. All collected individuals were found in ice above relatively shallow waters (60–100 m depth) suggesting bentho-sympagic coupling.     

Sympagic macro-fauna from multiyear sea-ice near Svalbard

Summary The object of the present investigation was to map the distribution and abundance of sympagic fauna (= ice fauna) (>350 m) within the perennial sea ice zone near Svalbard and to study relations between the sympagic fauna and the age and history of its ice substrate. The sampling took place in July/August 1986 and September 1988 using SCUBA-operated sampling gear (suction samplers, plankton nets with especially designed frames for sampling at the sea-ice/seawater interface, and underwater cameras). The amphipods Apherusa glacialis, Onisimus sp., and Gammarus wilkitzkii were the most conspicuous sympagic species both years. Scattered individuals of the amphipods Gammaracanthus loricatus, Weyprechtia penguis and the polychaete Harmathoinae indet. were also recorded. A. glacialis was the most numerous and contributed near 65% of the collected specimens in both years, with a maximum density exceeding 2000 individuals/m². However, G. wilkitzkii was on average larger, and contributed most to the biomass (1986: 80%; 1988: 77%). The average biomass of sympagic fauna in 1986 and 1988 was estimated to be 4.7 g/m² and 8.3 g/m² respectively. Biomass values reported here are ten to hundred times higher than what is found within the seasonal sea ice zone. Autochthonous sympagic species, like A. glacialis, Onisimus sp. and G. wilkitzkii, have a permanent association with ice. The seasonal sea ice zone will thus have to be recolonized every year resulting in lower densities compared to multiyear ice. It is suggested that the speed of the ice leaving the Polar Basin through the Fram Strait is too high for the sympagic fauna to remain its position in the sea ice zone. The result is an annual loss in the order of 7^*10⁵ tons of sympagic fauna from the perennial sea ice zone.     

On the distribution of sympagic macro-fauna in the seasonally ice covered Barents Sea

Summary The sympagic fauna (= sea ice fauna) of the Barents Sea was investigated on nine (2–5 weeks) cruises in the period 1983–1988. The amphipods Apherusa glacialis, Onisimus sp. and Gammarus wilkitzkii were the most conspicuous and abundant invertebrate species associated with the ice. Mean biomass-values ranged from 0 to about 2 g/m², and were less than 0.001 g/m² in five of the nine cruises. Apherusa glacialis, Onisimus sp. and Gammarus wilkitzkii are regarded as autochthonous sympagic species, and as such dependent upon a permanent ice cover. Allochthonous sympagic species were few and without significant importance. The variations in species abundance and biomass were believed to be closely connected to the age and drift-history of the ice. The density of the sympagic fauna increases as a result of increasing influx of ice into the Barents Sea from the Arctic Ocean. Density of the sympagic fauna were believed to increase with decreasing distance to the multiyear/first-year ice boundary. A. glacialis was found further away from the spreading centers and in higher numbers than the other sympagic species due to its greater motility. No day-night variations in the occurrences of the sympagic fauna were recorded.     

Under-ice distribution of polar cod <Emphasis Type="Italic">Boreogadus saida</Emphasis> in the central Arctic Ocean and their association with sea-ice habitat properties

In the Arctic Ocean, sea-ice habitats are undergoing rapid environmental change. Polar cod (Boreogadus saida) is the most abundant fish known to reside under the pack-ice. The under-ice distribution, association with sea-ice habitat properties and origins of polar cod in the central Arctic Ocean, however, are largely unknown. During the RV Polarstern expedition ARK XXVII/3 in the Eurasian Basin in 2012, we used for the first time in Arctic waters a Surface and Under Ice Trawl with an integrated bio-environmental sensor array. Polar cod was ubiquitous throughout the Eurasian Basin with a median abundance of 5000 ind. km^?2. The under-ice population consisted of young specimens with a total length between 52 and 140 mm, dominated by 1-year-old fish. Higher fish abundance was associated with thicker ice, higher ice coverage and lower surface salinity, or with higher densities of the ice-amphipod Apherusa glacialis. The fish were in good condition and well fed according to various indices. Back-tracking of the sea-ice indicated that sea-ice sampled in the Amundsen Basin originated from the Laptev Sea coast, while sea-ice sampled in the Nansen Basin originated from the Kara Sea. Assuming that fish were following the ice drift, this suggests that under-ice polar cod distribution in the Eurasian Basin is dependent on the coastal populations where the sea-ice originates. The omnipresence of polar cod in the Eurasian Basin, in a good body condition, suggests that the central Arctic under-ice habitats may constitute a favourable environment for this species survival, a potential vector of genetic exchange and a recruitment source for coastal populations around the Arctic Ocean.     

In-situ observations on the distribution and behavior of amphipods and Arctic cod (Boreogadus saida) under the sea ice of the High Arctic Canada Basin

The occurrence and behavior of sympagic amphipods and Arctic cod (Boreogadus saida) were studied in the High Arctic Canada Basin by diving under the ice at seven stations in summer 2002. Still images of video-transects were used to obtain animal abundances and information on the structure of the ice environment. Mean amphipod abundances for the stations varied between 1 and 23 individuals m^–2, with an increase towards the western part of the basin. The standard deviation within the 31–51 images analyzed per station was small (<1 individual m^–2). Gammarus wilkitzkii was found in low abundances, often hiding in small ice gaps. Small amphipods (Onisimus spp., Apherusa glacialis, and juveniles of all species) tended to move freely along the bottom of the floes. B. saida occurred in narrow wedges of seawater along the edges of melting ice floes at three stations in water depths of 10–50 cm and was never found under the ice. The fish occurred in schools of 1–28 per wedge. Fish were inactive and did not escape the approaching diver. Resting in the wedges may be a strategy to reduce energetic requirements and avoid predators.We dedicate this publication to Professor Dr. J. Lenz (Kiel University).     

New insights from old eggs – the shape and thickness of Great Auk Pinguinus impennis eggs

We compared the shape and eggshell thickness of Great Auk Pinguinus impennis eggs with those of its closest relatives, the Razorbill Alca torda, Common Guillemot Uria aalge and Brünnich's Guillemot Uria lomvia, in order to gain additional insights into the breeding biology of the extinct Great Auk. The egg of the Great Auk was most similar in shape to that of Brünnich's Guillemot. The absolute thickness of the Great Auk eggshell was greater than that of the Common Guillemot and Razorbill egg, which is as expected given its greater size, but the relative shell thickness at the equator and pointed end (compared with the blunt end) was more similar to that of the Common Guillemot. On the basis of these and other results we suggest that Great Auk incubated in an upright posture in open habitat with little or no nest, where its pyriform egg shape provided stability and allowed safe manoeuvrability during incubation. On the basis of a recent phylogeny of the Alcidae, we speculate that a single brood patch, a pyriform egg and upright incubation posture, as in the Great Auk and the two Uria guillemots, is the ancestral state, and that the Razorbill – the Great Auk's closest relative – secondarily evolved two brood patches and an elliptical egg as adaptations for horizontal incubation, which provides flexibility in incubation site selection, allowing breeding in enclosed spaces such as crevices, burrows or under boulders, as well as on open ledges.     

Surface-associated diatoms from marine habitats at Cape Evans,Antarctica, including the first record of living <Emphasis Type="Italic">Eunotogramma marginopunctatum</Emphasis>

The diversity of diatom communities found epiphytically on red macroalgae collected under the sea ice at Cape Evans, Antarctica, was studied microscopically. Comparison of communities along a depth profile showed that species diversity decreased as depth below sea ice increased. Dominant taxa also changed with depth, with Cocconeis fasciolata dominant at 10 and 15 m, Porosira glacialis at 20 m and Eunotogramma marginopunctatum at 25 m. Epiphytic communities were also compared to sympagic and sediment-associated communities collected from a single depth. Species richness was greatest for communities associated with surface sediments while the poorest was found in sympagic, or sea-ice, communities. E. marginopunctatum, previously only described from fossil material, was found associated epiphytically on macroalgae from 20 to 25 m below sea ice.     

Seasonal differences in at-sea activity of seabirds underline high energetic demands during the breeding period

We assessed seasonal differences in at-sea activity of Lesser Black-backed Gulls Larus fuscus, Black-legged Kittiwakes Rissa tridactyla and Common Guillemots Uria aalge in the south-eastern North Sea. The three species correspond to different ecological groups, with Lesser Black-backed Gulls representing omnivorous generalists, Kittiwakes representing surface-feeding pelagic seabirds, and Guillemots representing pursuit-diving pelagic seabirds. Using data from aerial surveys, we differentiated between active (flying or scavenging at fishing vessels) and inactive behaviour (swimming). We estimated the activity budgets of all three species for the different seasons and tested for differences in activity between different seasons. All species exhibited significant seasonal differences in activity, with the highest levels of activity observed during the breeding season. Numbers of flying auks were, however, exceptionally low in autumn due to moult and guarding of not-yet fledged chicks at sea. Our results underline the high energetic demands of the breeding season that lead to increased foraging and travelling activity.     

Flight speeds of two seabirds: a test of Norberg's hypothesis

Norberg suggested that birds should increase their flight speed when rearing chicks in order to maximize chick energy intake by reducing commuting time. We measured the incubation and chick-rearing flight speeds of a medium-range (Brünnich's Guillemot Uria lomvia ) and long-range (Northern Fulmar Fulmarus glacialis ) forager near the Prince Leopold Island colony, Nunavut, Canada. The mean flight speed for the long-range forager was significantly higher during chick-rearing than during incubation. The medium-range forager showed no difference in mean flight speed during the two periods. We suggest that because petrels fly close to their minimum power velocity and have a low wing-loading, whereas alcids fly close to their maximum range velocity and have a high wing-loading, petrels have a greater ability than alcids to alter their flight speed according to changes in the demands of different breeding stages. Consequently, whereas Northern Fulmars adapt to the additional cost of chick-rearing partially by altering flight speed, Brünnich's Guillemots can only do so by reducing mass.     

Vertical distribution of sympagic meiofauna in sea ice in the Canadian Beaufort Sea

This is the first study to determine vertical distribution patterns of sympagic meiofauna, including metazoans, protozoans and eggs >20 μm, in the Amundsen Gulf (southeastern Beaufort Sea, Arctic). Full sea-ice cores were sampled from mid of March to end of May 2008 (Circumpolar Flaw Lead system study). Investigations were performed on first-year ice from three pack- and three fast-ice stations. Additionally, 5-cm bottom-ice sections were sampled at 13 pack-ice and 5 fast-ice stations. The metazoan community was composed of nematodes, rotifers, copepods, copepod nauplii, platyhelminthes and a few rare taxa such as mollusks, cnidarians and nemerteans. High numbers of eggs, between 50 and 2,188 eggs L⁻¹, particularly of nematodes and copepods, were present in the ice. Investigations revealed also eggs of the pelagic species Calanus hyperboreus and Sagitta spp. within the ice, so that further research is needed to clarify whether more organisms than expected might use this habitat as a reproduction ground. Many different morphotypes of protozoans were observed in the samples, especially ciliates of the order Euplotida. The highest abundance was always found in the lowermost 5 cm of the ice cores, nevertheless sympagic meiofauna was not restricted to that part of the ice. Integrated meiofauna abundance ranged between 41 and 4,738 × 10² Ind. m⁻² and was highest in the fast ice in early May. Differences between pack and fast ice in terms of integrated meiofauna communities and vertical distribution were not significant, while the analysis of the bottom-ice sections indicated both a temporal development and ice-type-specific differences.     

A simple molecular approach to distinguish between two Arctic gadoid fishes <Emphasis Type="Italic">Arctogadus glacialis</Emphasis> (Peters, 1874) and <Emphasis Type="Italic">Boreogadus saida</Emphasis> (Lepechin, 1774)

Correct phenotypical identification of and discrimination between Boreogadus saida and Arctogadus glacialis is challenging, especially for larvae and young fish. We propose the use of a single microsatellite genetic marker, Gmo8, to distinguish between the two gadoid fish species. Amplified allele frequencies differed considerably, with B. saida (n = 97) being almost exclusively monomorphic at this locus, whereas A. glacialis (n = 136) is highly polymorphic. There was a clear separation between the amplified allele ranges for the two species. The species specific properties of Gmo8 enables the use of this marker to distinguish between B. saida and A. glacialis.     

Diving behaviour of Black Guillemots Cepphus grylle in the Pentland Firth,UK: potential for interactions with tidal stream energy developments

Capsule Black Guillemots Cepphus grylle were recorded diving in the Pentland Firth to an average depth of 32 metres and an overall maximal depth of 43 metres. The majority (88%) of dives were benthic with a median dive duration of 95 seconds, and a maximal dive duration of 131 seconds. The results provide empirical evidence that Black Guillemots use depths within the water column at which tidal turbines are likely to operate. Although limited, our data suggest the potential for interactions between Black Guillemots and marine renewable energy devices.     

An application of optimal diving models to diving behaviour of Brünnich's guillemots

Although theoretical models predict that the quality of foraging patches has little effect on optimal dive time with increasing depth, many empirical studies show that dive time at a given depth may vary. We developed a model that incorporated patch quality as a parameter of energy intake as a nonlinear function of time, and applied it to the diving behaviour of Brünnich's guillemots, Uria lomvia. The model indicated that optimal dive time can vary widely depending on the parameter. It also explained the convergence of observed dive times with travel time. Assuming the birds dived optimally, this parameter can be estimated from travel time and dive time for each dive. Foraging patches with larger estimated parameter values were favoured by the birds, suggesting that the parameter indicated patch quality. We used this parameter to test an optimal patch use model in divers. The results indicate that Brünnich's guillemots adjust their diving behaviour adaptively depending on patch quality, and that the optimal diving model is valid for prediction of observed dive patterns if patch quality is incorporated appropriately. Copyright 2002 The Association for the Study of Animal Behaviour. Published by Elsevier Science Ltd. All rights reserved.     

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.     

Kleptoparasitic interactions of Arctic Skuas Stercorarius parasiticus and Black Guillemots Cepphus grylle in northeastern Hudson Bay, Canada

Kleptoparasitic behaviour of Arctic Skuas was studied at a Black Guillemot colony in northeastern Hudson Bay, Canada. Skuas procured fish in 7-2% and 19-9% of attacks, and Black Guillemots lost fish in 7-2% and 21-2% of chases in 1982 and 1983, respectively. In 1983 chase outcome was related to distance between birds at the end of a chase and number of skuas in pursuit. The skuas' foraging efficiency (energy intake/energy expended during hunting) was estimated at 45 for 1982 and 5–8 for 1983, and total daily energy expenditure was estimated at 2–4 and 2-1 × b.m.r. for these years. Energetic calculations suggested that skuas obtained sufficient food to meet their daily needs in 3–6 h of hunting in 1982 and 2–7 h in 1983. The absence of intensive kleptoparasitic activity by breeding skuas in North America is attributed to the paucity of colonies suitable for both feeding (due to available host species and colony topography) and nesting (due to presence of arctic foxes and/or polar bears).     

Diet of black guillemots and northern fulmars breeding beside a High Arctic polynya

In the Arctic, the availability of early-season open water (shore leads, polynyas) is a key factor influencing annual reproduction by marine birds, as these relatively productive locations provide the only sites where migrating and breeding birds can feed. We examined the diet of two marine birds, black guillemots (Cepphus grylle) and northern fulmars (Fulmarus glacialis), feeding in the Hell Gate–Cardigan Strait Polynya in the Canadian High Arctic during the breeding seasons of 1980–1984. There was little overlap in prey items consumed between these two species, except that both relied on similar sympagic amphipods. Guillemots fed principally on benthic prey, and compared to earlier High Arctic studies, they consumed proportionally more benthic fish, mysids and decapods. Fulmars appeared to feed on swarms of pelagic or surface prey, and in this polynya they consumed proportionally fewer fish and more polychaetes than has been reported for fulmars at other High Arctic locations. Given that climate change may alter marine food webs by changing the types and availability of prey, our results set a baseline of dietary information for these predators against which future conditions can be compared.