Summer diet of seabirds feeding in sea-ice-covered waters near Svalbard

Summary We describe the summer diets of four seabird species. Kittiwake (Rissa tridactyla), Little Auk (Alle alle), Black Guillemot (Cepphus grylle) and Brünnich's Guillemot (Uria lomvia) collected in sea-ice-covered waters near Svalbard. Birds collected in an area filled with young sea-ice, within the seasonal sea-ice zone, were compared with birds collected from the perennial sea-ice zone dominated by multiyear ice. Pelagic Crustacea and fish dominated the diet of birds feeding in young ice, while sympagic Crustacea and fish were most important in the diet of birds feeding in multiyear ice. Boreogadus saida was the most important fish food item. B. Saida was present in the ice in both areas, while sympagic Crustacea were lacking in the area filled with young ice. Important food items in young ice were B. saida (Black Guillemots and Kittiwakes), Cajanus sp. (Little Auks) and Pandalus borealis (Brünnich's Guillemot). B. saida (Black Guillemots, Kittiwakes, Brünnich's Guillemots), Gammarus wilkitzkii (Brünnich's Guillemot, Black Guillemot) and Apherusa glacialis (Little Auk) were most important in multiyear ice. In general, Black Guillemots and Kittiwakes fed on fish, whereas Brünnich's Guillemots fed on the larger, and Little Auks on the smaller, Crustacea. The importance of sympagic species in the diet of seabirds is thought to be closely related to the age and history of the ice in the feeding area.     

Variations in Black Guillemot Cepphus grylle natal dispersal and philopatry across Britain and Ireland

Capsule: Using ring recovery records collected in Britain and Ireland from 1935 to 2015, we investigated philopatry and dispersal in Black Guillemots Cepphus grylle ringed as nestlings and recovered at breeding age during the breeding season. Levels of philopatry and dispersal distance varied between colonies, and were significantly related to latitude, possibly due to differences in ecology between populations. However, an increase in ringing effort is required to allow robust comparisons of these behaviours between colonies.     

Seasonal and annual variation in the diving behaviour of Hutton's shearwater (Puffinus huttoni)

ABSTRACT

With the development and implementation of tracking technology, we are now able to monitor the foraging behaviour of seabirds while at sea. Time-Depth Recorders (TDRs) were fitted to Hutton's shearwaters (Puffinus huttoni), an endangered endemic New Zealand species, to measure how diving behaviour varies over the breeding cycle. Hutton's shearwaters (～350?g) dive up to 339 times per day (average 68.8) at depths to 35?m (average 5.6?m), and for periods up to 60?s (average 19.2?s). Incubating birds dived deeper than birds feeding chicks, and a significant difference in diving depth and dive duration were detected at different times of the day. Neither dive frequency nor dive duration differed significantly between years, but there was some annual variation in dive depths. The temporal variation we observed in the diving behaviour of Hutton's shearwaters suggests they are likely to exploit different types of pelagic prey at different stages in their breeding cycle. With on-going changes in the marine environment, monitoring changes in feeding behaviour using TDRs may provide a way to assess environmental change and improve the conservation of this species.     

Foraging behaviour of Razorbills Alca torda during chick-rearing at the largest colony in the Baltic Sea

Capsule: Foraging behaviour in the Razorbill Alca torda during breeding was similar to that found elsewhere, aside from dive shape.

Aims: To investigate the foraging behaviour of Razorbills during the breeding season at the largest colony in the central Baltic Sea.

Methods: A combination of global positioning system (GPS) and time-depth recorder (TDR) devices were used on Razorbills breeding on the island of Stora Karlsö, Baltic Sea, during the chick-rearing period.

Results: Five GPS tracks and nine TDR logs were retrieved from 12 Razorbills, and 7399 dives were analysed. Razorbills foraged south and southwest of the colony. Maximum and mean (±sd) foraging range from the colony was 72.7?km and 13.1?±?13.5?km, respectively. Mean dive depth (15.3?±?2.4?m) and duration (53.1?±?8.5?s) were similar to those of a more southern Baltic Sea Razorbill colony. Dive depth had a bimodal distribution, with 70% of dives deeper than 10?m and 30% shallower than 10?m. There was a clear diel foraging pattern with 89% of dives occurring during daytime and a higher proportion of shallow dives at night. Unexpectedly, dives were primarily U-shaped. The Razorbills spent 31% of their overall time activity budget flying or diving.

Conclusion: Aside from dive shape, foraging behaviour was consistent with that reported at other colonies of Razorbills. Inconsistency in dive shape may be due to a bimodal foraging strategy, local prey behaviour or competition with the Common Guillemot Uria aalge.     

Artifacts arising from sampling interval in dive depth studies of marine endotherms

Most depth recorders used to study the diving behaviour of polar marine endotherms record depth data at specific time intervals. The length of recording interval can have potentially profound implications for the interpretation of the data. We used data acquired on the diving behaviour of king penguins, Aptenodytes patagonicus, to examine the validity of various analyses routinely conducted on depth data. In our experiments, increasing the sampling interval led to an underestimation of the number of dives performed, an overestimation in mean dive duration and substantial changes in the form of the dive profile. Our analysis indicates that depth data should be recorded at a minimum rate corresponding to 10% of the total dive duration and that conventional dive profile categorization may be inappropriate. Alternatives that are less subjective, and based on curve fits of dive depth versus time, are proposed.     

Finding your mate in a seabird colony: contrasting strategies of the Guillemot Uria aalge and King Penguin Aptenodytes patagonicus

Capsule King Penguins recognize their mates by voice, but Guillemots do not need acoustic cues even though their calls show individual variation.

Aims To determine whether the structure of Guillemot calls could allow individual recognition, as with King Penguin, and whether acoustic cues are used to locate mates among a dense mass of conspecifics at a colony.

Methods Observations were made on breeding Guillemots and King Penguins. Calls made by birds returning to their mates were recorded, the signals digitized and the calls analysed. Calls were later played back to the mates of the birds concerned and the effects noted on both them and their neighbours.

Results Both Guillemots and King Penguins emitted calls on return to the breeding site which contained individual signatures and were therefore potentially usable for mate recognition. In King Penguins, auditory recognition was essential for finding a mate, whereas in Guillemots most of the arriving birds located their mate in a dense crowd of conspecifics without the help of acoustic signals. Guillemots could differentiate neighbours from strangers without auditory cues.

Conclusion Calls are essential for the successful identification of mates by King Penguins but not by Guillemots.     

Dive bout organization in the Chinstrap penguin at seal island, antarctica

Diving behavior of 2 breeding Chinstrap penguins (Pygoscelis antarctica) was studied focusing first and primarily on dive bouts rather than dives themselves. Analysis of dive bout organization revealed (1) though there are differences between solitary dives and dive bouts in dive duration and dive depth, the first dives of dive bouts do not differ from solitary dives in the dive parameters, (2) mean dive duration during bout correlates positively to both mean dive depth during bout and mean surface interval during bout, while number of dives during bout negatively correlates to both cost (consumed energy) and duration of a dive cycle during bout. These findings suggest the following possibilities on foraging behavior of penguins: (1) their decision to repeat diving depends on the result of the first dive at a site, and the first dives of bouts would tend to be searching or evaluating dives though they would be also successful foraging dives, (2) they repeat diving at a foraging patch until foraging efficiency decrease to a threshold of diminishing returns.     

Foraging behavior of Adélie penguins during incubation period in Lützow-Holm Bay

The diving behavior of Adélie penguins Pygoscelis adeliae was investigated using time–depth recorders during the incubation period in the fast sea-ice area of Lützow-Holm Bay, Antarctica. Dive profiles and activity/time allocation suggested that penguins were obligated to walk on the fast-ice for 90–100 km until a polynya, which they used as an access to the pack-ice zone. Dive depth did not differ between males and females, though males’ dive duration was longer than that of females. Dive depth was slightly shallower and dive duration was shorter during the incubation than during the chick-rearing phase. Birds dove throughout the day, although less frequently around midnight, and there was no clear diel change in dive depth. This daily dive pattern during incubation period was similar to that previously observed during the chick-rearing period in a fast sea-ice area, but differed from that observed in sea-ice-free area. Variations in diving behavior resulted from different environmental conditions, such as foraging area with different sea-ice condition, as well as from different life history strategies.     

Dive types and circadian behaviour patterns of Saimaa ringed seals<Emphasis Type="Italic">Phoca hispida saimensis</Emphasis> during the open-water season

Diving and circadian behaviour patterns of 7 free-ranging Saimaa ringed sealsPhoca hispida saimensis Nordquist, 1899 were examined by VHF-radiotelemetry during open-water seasons between May and November in Lake Saimaa, eastern Finland. The mean recorded dive duration ranged from 2.8 to 6.5 min, with a maximum of 21 min. The mean dive depth ranged from 9.8 to 15.7 m, with maximum of 39.6 m. The maximum dive depth of each seal was limited by water depth in the study area. The dive depths were positively correlated with dive duration and body mass of the seal. Five different dive types were defined, as based on their depth-time characteristics, each falling into one of the three functional categories: travelling, feeding, and resting. Long duration diving bouts occurred mostly at night and were presumed to be resting dives. Saimaa ringed seals exhibited a circadian pattern of haul-out behaviour that shifted seasonally. During molting (May–June) the seals hauled-out both day and night, but later in summer haul-out was more frequent at night.     

Buoyancy control in diving behavior of the loggerhead turtle,Caretta caretta

Neutral buoyancy at the stationary depth is advantageous for diving animals. The adjustment of the air inspiration before diving can be a mechanism of buoyancy control for diving animals with lungs. The stationary depth of neutral buoyancy becomes deeper with larger inspiration. Our aim was to examine whether the loggerhead sea turtle,Caretta caretta regulates the buoyancy to be neutral at the stationary depth of the dive. During an internesting period of the breeding season, we recorded the diving pattern of an adult female using a time-depth recorder and a time-swim distance recorder. The dives were classified into four types (Types 1 to 4) based on the time-depth profile. Types-3 and 4 (66% of the total dive duration) have three phases in each dive: (1) first descent, (2) gradual ascent (stationary period), and (3) final ascent. In the gradual ascent phase, the turtle stayed at a certain depth without swimming. This means that the turtle was neutrally buoyant during the gradual ascent phase. The depth of the gradual ascent phase was positively correlated with the dive duration, supporting the hypothesis that neutral buoyancy of the loggerhead turtle is achieved by the air in their lungs.     

Diet and diving behaviour of European Storm Petrels Hydrobates pelagicus in the Mediterranean (ssp. melitensis)

Capsule Unlike Atlantic populations, which feed on krill, Mediterranean populations feed mainly on pelagic fish Gymnammodites cicerellus.

Aims To determine the diet and dive depth of the Mediterranean subspecies of European Storm Petrels Hydrobates pelagicus melitensis.

Methods Analysis of regurgitates of adults arriving at the colony for chick feeding and by determination of dives depth using the capillary tube method.

Results The main prey is Gymnammodites cicerellus, a pelagic fish. Storm Petrels dive for their prey and can reach up to 5 m in depth. They also make short foraging trips just outside the colony where they capture Opossum Shrimps Misydacea.

Conclusions European Storm Petrels in the Mediterranean exploit pelagic fish which are taken by diving. This contrasts with the Atlantic populations which feed mainly on krill. Mediterranean birds also feed on Opossum Shrimps Mysidacea during short foraging trips made at night just outside the colony. Differences in diet between long and short foraging trips may be because adults have to forage for both themselves and their chicks.     

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.     

The effects of quantity and quality of prey on population fluctuations in three seabird species

Capsule Population sizes of Common Guillemots Uria aalge, Razorbills Alca torda and Lesser Black‐backed Gulls Larus fuscus were associated with prey abundance but not prey quality.

Aims To examine how the abundance and quality of prey fish affects seabird population size and to test the ‘junk‐food’ or nutritional stress hypothesis.

Methods Analysis of long‐term seabird population size data and Sprat Sprattus sprattus biomass and age‐related weight data using a correlative approach.

Results De‐trended seabird and Sprat population data showed that the abundance of Sprat, the main prey species, was associated with the abundance of seabirds, while no effect of age‐related size of prey on seabird population size was found.

Conclusion As the Sprat population increased so did the seabird populations, regardless of decreases in ‘quality’ of Sprats, implying that more prey fish simply seem to mean more food in this marine ecosystem. No support for the ‘junk‐food’ hypothesis was found and the results contradict suggestions from earlier studies that prey quality is important to top‐predators in the Baltic Sea.     

Diving behavior in a free‐living,semi‐aquatic herbivore,the Eurasian beaver Castor fiber

Semi‐aquatic mammals have secondarily returned to the aquatic environment, although they spend a major part of their life operating in air. Moving both on land, as well as in, and under water is challenging because such species are considered to be imperfectly adapted to both environments. We deployed accelerometers combined with a depth sensor to study the diving behavior of 12 free‐living Eurasian beavers Castor fiber in southeast Norway between 2009 and 2011 to examine the extent to which beavers conformed with mass‐dependent dive capacities, expecting them to be poorer than wholly aquatic species. Dives were generally shallow (<1 m) and of short duration (<30 s), suggesting that the majority of dives were aerobic. Dive parameters such as maximum diving depth, dive duration, and bottom phase duration were related to the effort during different dive phases and the maximum depth reached. During the descent, mean vectorial dynamic body acceleration (VeDBA—a proxy for movement power) was highest near the surface, probably due to increased upthrust linked to fur‐ and lung‐associated air. Inconsistently though, mean VeDBA underwater was highest during the ascent when this air would be expected to help drive the animals back to the surface. Higher movement costs during ascents may arise from transporting materials up, the air bubbling out of the fur, and/or the animals’ exhaling during the bottom phase of the dive. In a manner similar to other homeotherms, beavers extended both dive and bottom phase durations with diving depth. Deeper dives tended to have a longer bottom phase, although its duration was shortened with increased VeDBA during the bottom phase. Water temperature did not affect diving behavior. Overall, the beavers’ dive profile (depth, duration) was similar to other semi‐aquatic freshwater divers. However, beavers dived for only 2.8% of their active time, presumably because they do not rely on diving for food acquisition.     

Tufted ducks Aythya fuligula do not control buoyancy during diving

Work against buoyancy during submergence is a large component of the energy costs for shallow diving ducks. For penguins, buoyancy is less of a problem, however they still seem to trade‐off levels of oxygen stores against the costs and benefits of buoyant force during descent and ascent. This trade‐off is presumably achieved by increasing air sac volume and hence pre‐dive buoyancy (B_pre) when diving deeper. Tufted ducks, Aythya fuligula, almost always dive with nearly full oxygen stores so these cannot be increased. However, the high natural buoyancy of tufted ducks guarantees a passive ascent, so they might be expected to decrease B_pre before particularly deep, long dives to reduce the energy costs of diving. Body heat lost to the water can also be a cause of substantial energy expenditure during a dive, both through dissipation to the ambient environment and through the heating of ingested food and water. Thus dive depth (d_d), duration and food type can influence how much heat energy is lost during a dive. The present study investigated the relationship between certain physiological and behavioural adjustments by tufted ducks to d_d and food type. Changes in B_pre, deep body temperature (T_b) and dive time budgeting of four ducks were measured when diving to two different depths (1.5 and 5.7 m), and for two types of food (mussels and mealworms). The hypothesis was that in tufted ducks, B_pre decreases as d_d increases. The ducks did not change B_pre in response to different diving depths, and thus the hypothesis was rejected. T_b was largely unaffected by dives to either depth. However, diving behaviour changed at the greater d_d, including an increase in dive duration and vertical descent speed. Behaviour also changed depending on the food type, including an increase in foraging duration and vertical descent speed when mussels were present. Behavioural changes seem to represent the major adjustment made by tufted ducks in response to changes in their diving environment.     

Water temperature sampling by foraging Brünnich's Guillemots with bird-borne data loggers

We describe the features of waters where seabirds were feeding by sampling vertical water temperature profiles with data loggers mounted on five Brünnich's Guillemots in Svalbard, Norway. The guillemots foraged in a cold water (−0.5–0.5°C SST (sea surface temperature)) by making 1.8 dive bouts in short trips (32–257 min duration) as well as in moderate (0.5–2.0°C SST) and warm waters (2.5–4.0°C SST) by making 6.0 dive bouts during long trips (411–688 min duration). Judging from outbound flying time (15.7–24.4 min), time between dive bouts (23.9–43.3 min) and water types, the birds probably fed in fjord or coastal waters during short trips and in both coastal and offshore waters during long trips. Water temperature and diving behaviour can be simultaneously recorded by small data loggers, which therefore will provide useful information on marine features and foraging activity of top predators.     

Effects of asexual reproduction on population structure of Sagartia elegans (Anthozoa: Actiniaria)

Sagartia elegans propagates asexually by pedal laceration. The effects of this method of reproduction on local population structure were investigated. Localized clones with discrete clonal boundaries occur. Expansion of individual clones may be extensive (tens of metres), but areas of uninhabitable substrate constitute an almost complete barrier to such dispersal. Short term, local recruitment is infrequent and appears to be dominated by pedal laceration but, despite the predominance of asexual reproduction, genotypic diversity is high, few populations being dominated by individual clones.     

Postrelease dive ability in rehabilitated harbor seals

The efficacy of seal rehabilitation is examined in a postrelease study of dive ability in harbor seal pups (Phoca vitulina) in the Wash, United Kingdom. Six rehabilitated seals were fitted with Sea Mammal Research Unit (SMRU) Argos Satellite Relay Data Logger tags and their individual dive behavior was monitored for an average of 122 d. The upper 90 percentile edge of dive behavior (dive duration [DD₉₀] and percentage of time at‐sea spent in a dive [PD₉₀]), in 7 d bins, was used as a proxy for physiological dive ability. The results are compared with data from five wild adult harbor seals. There was no statistically significant difference between (1) the mean track duration of rehabilitated seals (126.20 ± 27.48 [SD] d) and adult seals (150.2 ± 24.62 d) (P= 0.108), indicating no evidence that short‐term survival was less in the rehabilitated group; (2) the mean mass‐scaled DD₉₀ of rehabilitated seals (3.95 ± 0.37 min) and adult seals (4.09 ± 0.55 min) (P= 0.632); and (3) the mean PD₉₀ of rehabilitated seals (81.62 ± 1.21%) and adult seals (81.48 ± 3.93%) (P= 0.943). These three results all suggest the success of the rehabilitation program in terms of short‐term survival and dive ability.     

Benefits of thermal acclimation in a tropical aquatic ectotherm,the Arafura filesnake, <Emphasis Type="Italic">Acrochordus arafurae</Emphasis>

The presumption that organisms benefit from thermal acclimation has been widely debated in the literature. The ability to thermally acclimate to offset temperature effects on physiological function is prevalent in ectotherms that are unable to thermoregulate year-round to maintain performance. In this study we examined the physiological and behavioural consequences of long-term exposure to different water temperatures in the aquatic snake Acrochordus arafurae. We hypothesised that long dives would benefit this species by reducing the likelihood of avian predation. To achieve longer dives at high temperatures, we predicted that thermal acclimation of A. arafurae would reduce metabolic rate and increase use of aquatic respiration. Acrochordus arafurae were held at 24 or 32°C for 3 months before dive duration and physiological factors were assessed (at both 24 and 32°C). Although filesnakes demonstrated thermal acclimation of metabolic rate, use of aquatic respiration was thermally independent and did not acclimate. Mean dive duration did not differ between the acclimation groups at either temperature; however, warm-acclimated animals increased maximum and modal dive duration, demonstrating a longer dive duration capacity. Our study established that A. arafurae is capable of thermal acclimation and this confers a benefit to the diving abilities of this snake.     

Assessment of wind‐farm and other bird casualties from carcasses found on a Northumbrian beach over an 11‐year period

Capsule For 3748 bird carcasses found on 4.7 km of shoreline, the main cause of death was starvation. Three percent of deaths were attributed to wind turbines.

Aims To assess the main mortality causes from bird bodies washed ashore near wind turbines built in 1993.

Methods Weekly searches were made for bird carcasses to ascertain causes of death. Experiments tested the efficiency of searches, longevity of carcasses, and effects of wind direction on deposition rates.

Results In total, 3748 bird carcasses were found, an average of 341 per year. Guillemots formed 24.3% of the total, Kittiwakes 9.7%, Herring Gulls 9.0%, Black‐headed Gulls 7.4%, Great Black‐backed Gulls 6.4% and Feral Pigeons 11.3%. Each year more carcasses were found in winter than in summer, with a nine‐fold variation between winters. About 28.1% of carcasses were classed as starved, and 23.3% as eaten at sea (predation or scavenging). Of human‐related causes, 3.3% were birds affected by fishing gear, 3.0% were oiled, 6.4% had died from collisions (including 3% with wind‐turbines), the rest from minor or unidentified causes. Small passerines were probably under‐represented.

Conclusion Allowing for bodies not found, the local wind‐farm probably killed 148.5–193.5 birds per year, or 16.5–21.5 birds per turbine per year (mainly large gulls).