Fatty acid and stable isotope characteristics of sea ice and pelagic particulate organic matter in the Bering Sea: tools for estimating sea ice algal contribution to Arctic food web production

We determined fatty acid (FA) profiles and carbon stable isotopic composition of individual FAs (δ¹³C_FA values) from sea ice particulate organic matter (i-POM) and pelagic POM (p-POM) in the Bering Sea during maximum ice extent, ice melt, and ice-free conditions in 2010. Based on FA biomarkers, differences in relative composition of diatoms, dinoflagellates, and bacteria were inferred for i-POM versus p-POM and for seasonal succession stages in p-POM. Proportions of diatom markers were higher in i-POM (16:4n-1, 6.6–8.7 %; 20:5n-3, 19.6–25.9 %) than in p-POM (16:4n-1, 1.2–4.0 %; 20:5n-3, 5.5–14.0 %). The dinoflagellate marker 22:6n-3/20:5n-3 was highest in p-POM. Bacterial FA concentration was higher in the bottom 1 cm of sea ice (14–245 μg L^?1) than in the water column (0.6–1.7 μg L^?1). Many i-POM δ¹³C_FA values were higher (up to ~10 ‰) than those of p-POM, and i-POM δ¹³C_FA values increased with day length. The higher i-POM δ¹³C_FA values are most likely related to the reduced dissolved inorganic carbon (DIC) availability within the semi-closed sea ice brine channel system. Based on a modified Rayleigh equation, the fraction of sea ice DIC fixed in i-POM ranged from 12 to 73 %, implying that carbon was not limiting for primary productivity in the sympagic habitat. These differences in FA composition and δ¹³C_FA values between i-POM and p-POM will aid efforts to track the proportional contribution of sea ice algal carbon to higher trophic levels in the Bering Sea and likely other Arctic seas.     

Summer diet of beluga whales inferred by fatty acid analysis of the eastern Beaufort Sea food web

Beluga whales (Delphinapterus leucas) are the most abundant odontocetes in Arctic waters and are thus thought to influence food web structure and function. The diet of the Beaufort Sea beluga population is not well known, partly due to the inherent difficulty of observing feeding behaviour in Arctic marine cetaceans. To determine which prey items are critical to the Beaufort Sea beluga diet we first examine and describe the Mackenzie Delta and Beaufort Sea food web using fatty acid analyses. Fatty acid profiles effectively partitioned prey items into groups associated with their habitat and feeding ecology. Next, the relative contribution of various prey items to beluga diet was investigated using fatty acids. Finally, beluga diet variability was examined as a function of body size, a known correlate of habitat use. Beluga appeared to feed predominantly on Arctic cod (Boreogadus saida) collected from near shore and offshore regions. Size related dietary differences suggested larger sized beluga preferred offshore Arctic cod given the shared high levels of long chain monounsaturates, whereas smaller sized beluga appeared to feed on prey in their near shore habitats that included near shore Arctic cod. The presence of Arctic cod groups in shallow near shore and deep offshore habitats may facilitate the behavioural segregation of beluga habitat use as it relates to their size and resource requirements. Given Arctic cod are a sea ice associated fish combined with the accelerated sea ice loss in this region, beluga whales may need to adapt to new dietary regimes.     

Fatty acid composition of the blubber in white whales (Delphinapterus leucas)

Fatty acid (FA) composition of the blubber in free-ranging white whales (Delphinapterus leucas) from Svalbard's waters was determined and compared with the fatty acid composition of potential prey species in an attempt to assess diet. This methodology is based on the common assumption that unique arrays of FAs found within groups of organisms are transferred, largely unaltered, up marine food chains and thus may be useful for assessment of diet composition. Complete-column blubber biopsies were sampled from white whales (n=7) during the summers of 1996 and 1997. All captured animals were adult males. FAs were extracted from 2–4 replicates taken from an area about 10 cm in front of the mid-dorsal ridge. FA data from a total of 12 potential prey species from the Svalbard area were compared to the white-whale blubber samples. Twenty-two FAs were consistently found in relative amounts >0.5% of the total FA composition in white whales. These FAs accounted for 94–96% of the total FAs present. The blubber was composed almost entirely of triacylglycerols. The major saturated FAs were 14:0 and 16:0; 16:1(n-7), 18:1(n-9) and 20:1(n-9) were the major monounsaturated FAs and 20:5(n-3) and 22:6(n-3) were the major polyunsaturated FAs. Sixteen of the 22 FAs consistently found in the white-whale blubber were also found in considerable amounts (>0.5% of total FAs) in most of the potential species. Principal Component Analysis run on these 16 FAs suggests that polar cod (Boreogadus saida) had the most similar FA composition to the white-whale blubber, followed by capelin (Mallotus villosus), the copepod Calanus hyperboreus and the shrimp Pandalus borealis. Accepted: 27 November 1999     

Resource partitioning between Pacific walruses and bearded seals in the Alaska Arctic and sub-Arctic

Climate-mediated changes in the phenology of Arctic sea ice and primary production may alter benthic food webs that sustain populations of Pacific walruses (Odobenus rosmarus divergens) and bearded seals (Erignathus barbatus). Interspecific resource competition could place an additional strain on ice-associated marine mammals already facing loss of sea ice habitat. Using fatty acid (FA) profiles, FA trophic markers, and FA stable carbon isotope analyses, we found that walruses and bearded seals partitioned food resources in 2009–2011. Interspecific differences in FA profiles were largely driven by variation in non-methylene FAs, which are markers of benthic invertebrate prey taxa, indicating varying consumption of specific benthic prey. We used Bayesian multi-source FA stable isotope mixing models to estimate the proportional contributions of particulate organic matter (POM) from sympagic (ice algal), pelagic, and benthic sources to these apex predators. Proportional contributions of FAs to walruses and bearded seals from benthic POM sources were high [44 (17–67)% and 62 (38–83)%, respectively] relative to other sources of POM. Walruses also obtained considerable contributions of FAs from pelagic POM sources [51 (32–73)%]. Comparison of δ¹³C values of algal FAs from walruses and bearded seals to those from benthic prey from different feeding groups from the Chukchi and Bering seas revealed that different trophic pathways sustained walruses and bearded seals. Our findings suggest that (1) resource partitioning may mitigate interspecific competition, and (2) climate change impacts on Arctic food webs may elicit species-specific responses in these high trophic level consumers.     

Fatty acids as trophic tracers in an experimental estuarine food chain: Tracer transfer

The transfer of fatty acid (FA) biomarkers was assessed by an experimental food chain comprising three trophic levels: leaves of the mangrove Avicennia marina, the grapsid crab Parasesarma erythodactyla, and the blue swimmer crab Portunus pelagicus. FA compositions for each trophic level were investigated through a feeding/starving regime designed to reveal the transfer of FAs along the food chain. Comparison of the FA profiles of the mangrove leaves, the tissues and faecal material of P. erythodactyla suggests that the crab, contrary to previous studies, lacks the necessary enzymes to incorporate some FAs in its diet. Long chain FAs were egested while polyunsaturated acids seemed to be efficiently assimilated. The polyunsaturated FAs 18:2ω6 and 18:3ω3 were identified as useful biomarkers of the mangrove leaves for tracing their transfer to the higher trophic levels. The contribution of these markers to the FA profiles of the crabs was investigated and it was found that both 18:2ω6 and 18:3ω3 could be successfully traced across the first trophic transfer. However, only 18:3ω3 demonstrated a clear second transfer into the tissues of P. pelagicus. Multivariate analysis of the FA profiles of the study organisms was found to be a potentially useful tool for demonstrating differences in diet within a species and also what FAs, and therefore dietary items, are responsible for those differences. MDS analysis of the FA profiles of faecal material from P. erythodactyla showed that this species provides an important ecological link in estuarine systems by providing a substrate for the colonisation of bacteria.     

Temporal Dynamics of Top Predators Interactions in the Barents Sea

The Barents Sea system is often depicted as a simple food web in terms of number of dominant feeding links. The most conspicuous feeding link is between the Northeast Arctic cod Gadus morhua, the world''s largest cod stock which is presently at a historical high level, and capelin Mallotus villosus. The system also holds diverse seabird and marine mammal communities. Previous diet studies may suggest that these top predators (cod, bird and sea mammals) compete for food particularly with respect to pelagic fish such as capelin and juvenile herring (Clupea harengus), and krill. In this paper we explored the diet of some Barents Sea top predators (cod, Black-legged kittiwake Rissa tridactyla, Common guillemot Uria aalge, and Minke whale Balaenoptera acutorostrata). We developed a GAM modelling approach to analyse the temporal variation diet composition within and between predators, to explore intra- and inter-specific interactions. The GAM models demonstrated that the seabird diet is temperature dependent while the diet of Minke whale and cod is prey dependent; Minke whale and cod diets depend on the abundance of herring and capelin, respectively. There was significant diet overlap between cod and Minke whale, and between kittiwake and guillemot. In general, the diet overlap between predators increased with changes in herring and krill abundances. The diet overlap models developed in this study may help to identify inter-specific interactions and their dynamics that potentially affect the stocks targeted by fisheries.     

Fatty acid profiles of algae mark the development and composition of harpacticoid copepods

1. The value of algal fatty acids (FA) as diet biomarkers for benthic harpacticoid copepods was investigated. A high proportion of 18:1ω9 and 18:2ω6 FA was observed in the lipid reserve fraction of copepods fed with cyanobacteria. In contrast, a high proportion of 16:1ω7 and ω3 FA (including eicosapentaenoic) was present in the lipid reserve fraction of copepods grown on diatoms. 2. Copepods that were grown on cyanobacteria showed reduced survival and took 26% more time to develop from the first copepodid stage to adult than copepods that were grown on diatoms. Copepods feeding on the cyanobacteria showed reduced FA content when compared with animals fed with diatoms. This reduction in FA content was more pronounced in the apolar lipid fraction (mainly reserve lipids) than in the polar (mainly structural) lipid fraction. 3. The FA profiles of algae were used to calculate a function discriminating between diatoms and cyanobacteria. This function was applied to the FA profiles in the reserve lipid fraction of copepods and correctly classified copepod diet. 16:1ω7, 18:2ω6 and 20:5ω3 were the most important FA in the discriminant function. The suitability of this chemometric method to infer copepod diet was further tested by using algal class FA data from literature to derive the discriminant functions. The correct classification of the diet when the functions were applied to FA composition of the copepod reserve lipids suggests that this method may be employed in trophic web studies. 18:3ω3, 18:1ω9 and 16:1ω7 were the most important FA in the functions discriminating diatoms, cyanobacteria and green algae. The identification and quantification of the whole suit of 16:1ω7, 18:1ω9, 18:2ω6, 18:3ω3 and 20:5ω3 in trophic web studies is therefore of paramount importance to infer diet origin of aquatic herbivores. 4. The FA profile of copepod polar lipids did not reflect that of the diet. The presence of long chain polyunsaturated FAs in the polar lipid fraction of copepods feeding on the cyanobacterium suggests that C18 FAs from the diet may be elongated and desaturated by the copepod. The ability to elongate and desaturated FAs may reduce the importance of some FAs as diet biomarkers while it may turn the copepods into valuable trophic intermediaries in transferring organic matter from microorganisms to higher trophic levels.     

Body regional distribution and stratification of fatty acids in the blubber of New Zealand sea lions: implications for diet predictions

Fatty acids (FAs) from blubber are often analysed to assess the diet of marine mammals. However, distribution of blubber FAs is not necessarily uniform along the body. It is therefore important to understand the deposition of dietary fat to be able to estimate the diet. We analysed the FA compositions of the thoracic ventral (T region) blubber of 28 New Zealand (NZ) sea lions Phocarctos hookeri by-caught by the southern arrow squid Nototodarus sloani fishery. Each blubber sample was divided into an inner and an outer layer. For 16 of these 28 animals, the pelvic dorsal (P) region was also sampled. The influence of body region and layer was statistically tested on the distribution of blubber FAs. We found minimal differences between the P and T regions (3 out of 29 FAs). The outer blubber layer was more concentrated in short-chain monounsaturated FAs, and less concentrated in saturated FAs, but the degree of stratification was small. Diet predictions from quantitative FA signature analysis (QFASA) applied on different body regions were similar. When applied to different blubber layers, QFASA gave some variation in the contribution of rattails (~25 % in outer blubber vs. ~12 % in inner blubber). Nonetheless, diet predicted from both layers was dominated by similar prey species: octopus, hoki and rattails. Hoki and rattails shared a similar ecological niche. Therefore, feeding ecology of NZ sea lions inferred from the inner or the outer blubber would lead to the same conclusions. In the case of NZ sea lions, the outer layer of blubber, if the only sample accessible, could be a useful tissue for diet inference from FAs.     

The diet of polar bears (Ursus maritimus) from Svalbard, Norway, inferred from scat analysis

Polar bears (Ursus maritimus) are heavily dependent on marine prey, in particular ice-associated seals, which they hunt on landfast sea ice or free-floating pack ice. Dramatic current (and predicted) losses of sea ice habitat make it increasingly important to gain more knowledge of the relative use by bears of all types of prey from the marine food web as well as from terrestrial sources. This study uses frequency of occurrence of food items in 119 polar bear scats sampled on the sea ice as well as on shore in coastal areas in the Svalbard Archipelago, mainly in spring, between 2003 and 2010 to explore the diet of bears in the region. Ringed seals (Pusa hispida) occurred in 62.2 % (CI 52.8–70.9 %) of the scat samples examined. Various terrestrial plants (32.8 %, CI 24.4–42.0 %) and marine algae (21.8 %, CI 14.8–30.4 %) also occurred frequently in the scats; the significance of this high occurrence of plants and algae is not clear. Bearded seals (Erignathus barbatus) and various bird species constituted only minor components of the diet, while Svalbard reindeer (Rangifer tarandus platyrhynchus) occurred in 9.2 % (CI 4.7–15.9 %) of the scats, indicating that this species may play a more important role than previously reported. The novel combination of genetic analyses of material in the fecal samples along with detailed exploration of the physical–structural properties of prey hairs and plant parts provided a much fuller picture of the diet of polar bears than would have been possible from observational studies of polar bear predation behavior alone. This approach may provide an important tool for monitoring the responses of polar bears to ongoing ecosystem changes that will result from continued warming in the Arctic.     

Changes in the fatty acid composition of symbiotic dinoflagellates from the hermatypic coral <Emphasis Type="Italic">Echinopora lamellosa</Emphasis> during adaptation to the irradiance level

The composition of fatty acids (FAs) of symbiotic dinoflagellates isolated from the hermatypic coral Echinoporal lamellosa adapted to the irradiance of 95, 30, 8, and 2% PAR was studied. Polar lipids and triacylglycerols (TAG) differed between them in FA composition. Polar lipids were enriched in unsaturated FAs, whereas TAG, in saturated FAs. Light exerted a substantial influence on the FA composition in both polar lipids and TAG. The elevation of irradiance resulted in the accumulation of 16:0 acid in both lipid groups and 16:1(n-7) acid in TAG. It seems likely that de novo synthesis of 16:0 acid occurred actively in the cells of symbiotic dinoflagellates in high light. Since these processes are energy-consuming ones, they utilize excessive energy. When light intensity declined, 18:4(n-3) and 20:5(n-3) acids accumulated in polar lipids, which was accompanied by the increase in the content of chlorophyll a in the cells of zooxanthellae, whereas the levels of 22:6(n-3) and 20:4(n-6) acids reduced. Although the relative content of particular FAs varied substantially in dependence of irradiance, the balance between the sum of saturated and unsaturated FAs changed insignificantly. We concluded that the role of photoadaptation could not be limited only to changes in the degree of lipid unsaturation and membrane fluidity. It is supposed that light-induced changes in the FA composition reflect the interrelation between photosynthesis and FA biosynthesis.     

Temporal dynamics of amino and fatty acid composition in the razor clam Ensis siliqua (Mollusca: Bivalvia)

Few studies have been conducted on the temporal dynamics of both amino acid (AA) and fatty acid (FA) profiles in marine bivalves. We investigated the seasonal variation of these compounds in the pod razor clam Ensis siliqua in relation to food availability, salinity, water temperature and reproductive cycle. AA content varied between 46.94 and 54.67 % dry weight (DW), and the AAs found in greater quantity were glutamic acid, glycine and aspartic acid. FA content varied between 34.02 and 87.94 mg g^?1 DW and the FAs found in greater quantity were 16:0 and 22:6n-3. Seasonal trends were observed for AAs and FAs. FAs increased with gametogenesis and decreased with spawning while AA content increased throughout spawning. The effect of increasing temperature and high food availability during the spawning season masked the loss of AAs resulting from gamete release. Still, a comparatively greater increase in the contents of glutamic acid and leucine with spawning indicate their possible involvement in a post-spawning gonad recovery mechanism. A post-spawning decrease in 14:0, 16:0, 16:1n-7, 18:1n-7 and 18:1n-9 is indicative of the importance of these FAs in bivalve eggs. An increase in 18:3n-3, 18:4n-3, 20:1n-9 and 20:2n-6 during gametogenesis suggests their involvement in oocyte maturation. The FA 22:4n-6, while increasing with spawning, appears to play a role in post-spawning gonad recovery. Salinity did not have an effect on the AA composition. None of the environmental parameters measured had an effect on FA composition.     

Dependency of Antarctic zooplankton species on ice algae‐produced carbon suggests a sea ice‐driven pelagic ecosystem during winter

How the abundant pelagic life of the Southern Ocean survives winter darkness, when the sea is covered by pack ice and phytoplankton production is nearly zero, is poorly understood. Ice‐associated (“sympagic”) microalgae could serve as a high‐quality carbon source during winter, but their significance in the food web is so far unquantified. To better understand the importance of ice algae‐produced carbon for the overwintering of Antarctic organisms, we investigated fatty acid (FA) and stable isotope compositions of 10 zooplankton species, and their potential sympagic and pelagic carbon sources. FA‐specific carbon stable isotope compositions were used in stable isotope mixing models to quantify the contribution of ice algae‐produced carbon (α_Ice) to the body carbon of each species. Mean α_Ice estimates ranged from 4% to 67%, with large variations between species and depending on the FA used for the modelling. Integrating the α_Ice estimates from all models, the sympagic amphipod Eusirus laticarpus was the most dependent on ice algal carbon (α_Ice: 54%–67%), and the salp Salpa thompsoni showed the least dependency on ice algal carbon (α_Ice: 8%–40%). Differences in α_Ice estimates between FAs associated with short‐term vs. long‐term lipid pools suggested an increasing importance of ice algal carbon for many species as the winter season progressed. In the abundant winter‐active copepod Calanus propinquus, mean α_Ice reached more than 50% in late winter. The trophic carbon flux from ice algae into this copepod was between 3 and 5 mg C m^?2 day^?1. This indicates that copepods and other ice‐dependent zooplankton species transfer significant amounts of carbon from ice algae into the pelagic system, where it fuels the food web, the biological carbon pump and elemental cycling. Understanding the role of ice algae‐produced carbon in these processes will be the key to predictions of the impact of future sea ice decline on Antarctic ecosystem functioning.     

Fatty acid composition of sprat (<Emphasis Type="Italic">Sprattus sprattus</Emphasis>) and herring (<Emphasis Type="Italic">Clupea harengus</Emphasis>) in the Baltic Sea as potential prey for salmon (<Emphasis Type="Italic">Salmo salar</Emphasis>)

Sprat (Sprattus sprattus) and small herring (Clupea harengus) are the dominant prey fish of Atlantic salmon (Salmo salar) in the Baltic Sea. If the fatty acid (FA) proportions of sprat and herring differ, the dietary history of ascending salmon could be determined from their FA profiles. Therefore, we investigated the FA composition of several age groups of whole sprat and small herring, caught from the three main feeding areas of salmon in autumn and spring. Oleic acid (18:1n-9) was the most prevalent FA in sprat and characteristic of this species. In herring, palmitic acid (16:0) was the most common FA, but herring lipid was characterized by n-6 polyunsaturated FAs, and moreover, by palmitoleic acid (16:1n-7) and vaccenic acid (18:1n-7). Due to the higher lipid content of sprat, the concentrations of all other FAs, excluding these, were higher in sprat than in herring. The concentration of docosahexaenoic acid (DHA, 22:6n-3) increased with an increase in the lipid content and was consequently highest in the youngest specimens, being in young sprat almost double that of young herring, and 2.6 times higher in the sprat biomass than in that of herring. As a result of a decrease in the DHA concentration with age, the ratio thiamine/DHA increased with respect to age in both species, and was lower in sprat than in herring. It is concluded that an abundance of DHA in the diet of salmon most likely increases oxidative stress because of the susceptibility of DHA to peroxidation, and thus decreases thiamine resources of fasting, prespawning salmon. Because the FA composition of sprat and herring differs, and the relative abundancies of prey fish differ between the feeding areas of salmon, the feeding area of ascending salmon can most probably be derived by comparing their FA profiles.     

Is the poleward expansion by Atlantic cod and haddock threatening native polar cod, <Emphasis Type="Italic">Boreogadus saida</Emphasis>?

During a recent period of increased influx of warm Atlantic water to the western coast of Svalbard, we have observed a northward expansion of boreal Atlantic cod (Gadus morhua) and haddock (Melanogrammus aeglefinus) into areas dominated by the native polar cod (Boreogadus saida). To determine the potential impact of new ecological interactions, we studied the diet of co-occurring juvenile gadoids in fjords, open water, and sea ice around Svalbard. We also reviewed the available literature on polar cod feeding in different habitats across the Arctic to determine whether region, habitat, or fish size may influence diet. Feeding by polar cod in the pelagic zone was size dependent, with small fish primarily consuming Calanus spp. and smaller copepods, with an increasing ration of Themisto spp. at larger sizes. In benthic habitats, diets were more varied and included considerably more unidentified material and sediment. Less than 40% dietary overlap was detected among the three species when they were found together. Stable isotope analyses indicated these patterns were representative of longer-term assimilation. The low interspecific dietary overlap suggests little direct competition. Future increases in abundance and the high predation potential of the boreal taxa, however, may impact the persistence of polar cod on some Arctic shelves.     

Temporal fatty acid dynamics of the octocoral Veretillum cynomorium

The objectives of the present work were to investigate the temporal variation in the fatty acid (FA) composition of the octocoral Veretillum cynomorium, examine the effects of reproduction and environmental factors on FA variation, and establish a chemotaxonomic identification for this species. Mean oocyte size-frequency distributions showed that the majority of the oocytes had an intermediate size (Group II) before spawning (April and June). The late-vitellogenic oocytes (Group III) became absent in August and October and, during this post-spawning period, oocytes were primarily of small size (Group I). Most of the major FA, 16:0, 18:0, 20:4n-6, 20:5n-3, and the tetracosapolyenoic fatty acid (TPA), 24:6n-3, varied significantly throughout the year (p < 0.01), with two peaks in August/October and February. The boost in early oogenesis, also associated with warmer temperatures, seemed to be responsible for the observed increase in FA content between June and August. The highest values of FA content were observed in February when intermediate oogenesis (Group II) was at its peak and there were considerable levels of available food in the environment. Also, the increase in food availability seemed to trigger the final stages of gametogenesis. The high quantity of 18:1n-7, odd-numbered and branched FAs, suggested the presence of a dynamic bacterial community in V. cynomorium, probably as an adaptive response to the lack of symbiotic microalgae. Although the presence of TPAs is the main feature distinguishing octocorals from other coral species, here we showed that there was no single FA clearly dominating the FA composition of V. cynomorium throughout the year. Instead, four main FAs share similar concentrations: 16:0, 20:4n-6, 20:5n-3 and 24:6n-3. The predominance of these four FAs combined with the higher amount of 24:6n-3 when compared to 24:5n-6 may serve as a chemotaxonomic feature to distinguish this octocoral species (or genus).     

Inter-annual variability in the proportional contribution of higher trophic levels to the diet of Pacific walruses

Pacific walruses (Odobenus rosmarus divergens) depend on Arctic sea ice as a resting and foraging platform; however, recent years have seen unprecedented seasonal reductions in ice extent. Previous researchers proposed that during unfavorable ice conditions, walruses might prey on other pinnipeds. To examine this hypothesis, we analyzed carbon and nitrogen stable isotope ratios of muscle from walruses (n = 155) sampled from the Bering and Chukchi seas during 2001–2010. We used a Bayesian stable isotope mixing model to examine the proportional contribution of higher trophic level prey (HTLP) (e.g., seals, seabirds) to walrus diets and extrapolated a tissue-specific turnover rate to compare diet of individuals over time. Mode HTLP across years was 19 % ± 8. Results indicate a significant decrease (P < 0.05) in the reliance on HTLP during 2008–2009 (mode HTLP 13 %), one of two sampling periods that experienced great seasonal loss of pan-Arctic sea ice (the other being 2007–2008 with mode HTLP of 23 %). We also reveal intra-annual fluctuations in the contribution of HTLP to the diet of a walrus sampled in 2011 with seal remains in its stomach through high-resolution sectioning along a whisker length. Our findings suggest that walruses forage opportunistically as a result of multiple environmental factors and that sea ice extent alone does not drive consumption of HTLP.     

Variability in the summer diets of juvenile polar cod (<Emphasis Type="Italic">Boreogadus saida</Emphasis>) in the northeastern Chukchi and western Beaufort Seas

Polar cod (Boreogadus saida) is an important link between top predators and lower trophic levels in high-latitude marine ecosystems. Previous findings describe differences in its diet throughout the western Arctic; however, the causes of this variation are not well known. This study examined the diets of juvenile polar cod collected via demersal trawling methods over three summers in the northeastern Chukchi Sea (2010–2012) and one summer in the western Beaufort Sea (2011) to determine the amount of variability explained by biological, spatial, and interannual factors. Prey were identified, measured for length, and aggregated by percent mean weight into taxonomically coarse prey categories for analysis. Within seas, variation in juvenile polar cod diet composition was significantly related to body size, latitude, longitude, depth, and interannual (Chukchi Sea only) factors. Canonical correspondence analysis indicated body size was the most important factor contributing to the total variance in juvenile polar cod diet in the Chukchi and Beaufort Seas. Body size-based diet differences between the Chukchi and Beaufort Seas were evaluated using non-metric multidimensional scaling. This method revealed that similar-sized polar cod consumed similar-sized prey in both seas, but their diets were more benthically influenced in the Chukchi Sea and more pelagically influenced in the Beaufort Sea. Juvenile polar cod diet compositions vary by body size and region of inhabitance throughout their distribution. Here, we show that body size was the primary factor explaining variation in the summer diet of juvenile polar cod within the Chukchi and Beaufort Seas.     

Tracing carbon flow in an arctic marine food web using fatty acid-stable isotope analysis

Global warming and the loss of sea ice threaten to alter patterns of productivity in arctic marine ecosystems because of a likely decline in primary productivity by sea ice algae. Estimates of the contribution of ice algae to total primary production range widely, from just 3 to >50%, and the importance of ice algae to higher trophic levels remains unknown. To help answer this question, we investigated a novel approach to food web studies by combining the two established methods of stable isotope analysis and fatty acid (FA) analysis--we determined the C isotopic composition of individual diatom FA and traced these biomarkers in consumers. Samples were collected near Barrow, Alaska and included ice algae, pelagic phytoplankton, zooplankton, fish, seabirds, pinnipeds and cetaceans. Ice algae and pelagic phytoplankton had distinctive overall FA signatures and clear differences in delta(13)C for two specific diatom FA biomarkers: 16:4n-1 (-24.0+/-2.4 and -30.7+/-0.8 per thousand, respectively) and 20:5n-3 (-18.3+/-2.0 and -26.9+/-0.7 per thousand, respectively). Nearly all delta(13)C values of these two FA in consumers fell between the two stable isotopic end members. A mass balance equation indicated that FA material derived from ice algae, compared to pelagic diatoms, averaged 71% (44-107%) in consumers based on delta(13)C values of 16:4n-1, but only 24% (0-61%) based on 20:5n-3. Our estimates derived from 16:4n-1, which is produced only by diatoms, probably best represented the contribution of ice algae relative to pelagic diatoms. However, many types of algae produce 20:5n-3, so the lower value derived from it likely represented a more realistic estimate of the proportion of ice algae material relative to all other types of phytoplankton. These preliminary results demonstrate the potential value of compound-specific isotope analysis of marine lipids to trace C flow through marine food webs and provide a foundation for future work.     

Spring fasting behavior in a marine apex predator provides an index of ecosystem productivity

The effects of declining Arctic sea ice on local ecosystem productivity are not well understood but have been shown to vary inter‐specifically, spatially, and temporally. Because marine mammals occupy upper trophic levels in Arctic food webs, they may be useful indicators for understanding variation in ecosystem productivity. Polar bears (Ursus maritimus) are apex predators that primarily consume benthic and pelagic‐feeding ice‐associated seals. As such, their productivity integrates sea ice conditions and the ecosystem supporting them. Declining sea ice availability has been linked to negative population effects for polar bears but does not fully explain observed population changes. We examined relationships between spring foraging success of polar bears and sea ice conditions, prey productivity, and general patterns of ecosystem productivity in the Beaufort and Chukchi Seas (CSs). Fasting status (≥7 days) was estimated using serum urea and creatinine levels of 1,448 samples collected from 1,177 adult and subadult bears across three subpopulations. Fasting increased in the Beaufort Sea between 1983–1999 and 2000–2016 and was related to an index of ringed seal body condition. This change was concurrent with declines in body condition of polar bears and observed changes in the diet, condition and/or reproduction of four other vertebrate consumers within the food chain. In contrast, fasting declined in CS polar bears between periods and was less common than in the two Beaufort Sea subpopulations consistent with studies demonstrating higher primary productivity and maintenance or improved body condition in polar bears, ringed seals, and bearded seals despite recent sea ice loss in this region. Consistency between regional and temporal variation in spring polar bear fasting and food web productivity suggests that polar bears may be a useful indicator species. Furthermore, our results suggest that spatial and temporal ecological variation is important in affecting upper trophic‐level productivity in these marine ecosystems.     

Fatty‐acid biomarkers and tissue‐specific turnover: validation from a controlled feeding study in juvenile Atlantic croaker Micropogonias undulatus

Fatty‐acid (FA) profiles of liver and muscle tissue from juvenile Atlantic croaker Micropogonias undulatus were examined over a 15 week diet‐switch experiment to establish calibration coefficients (CC) and improve understanding of consumer–diet relationships for field applications. Essential FAs [docosahexaenoic acid (DHA), 22:6n‐3 and eicosapentaenoic acid (EPA) , 20:5n‐3] decreased and 18:2n‐6 increased in tissues of M. undulatus fed diets with increasing proportions of terrestrial v. marine lipid sources. Non‐linear models used to estimate the incorporation rate and days to saturation of per cent 18:2n‐6 in tissues showed that livers incorporated 18:2n‐6 faster than muscle, but the proportions of 18:2n‐6 in muscle were higher. CCs were established to determine proportions of FA deposition in tissues relative to diet. Many CCs were consistent amongst diet treatments, despite growth and dietary differences. The CCs can be used to discern FA modification and retention within tissues and as tools for future quantitative estimates of diet histories. Incorporation rates and CCs of 18:2n‐6 were applied to a sub‐set of field samples of wild M. undulatus to understand habitat use and feeding ecology. Altogether, these results suggest that FAs provide a time‐integrated measure of diet in aquatic food webs and are affected by tissue type, growth rate and the influence of mixed diets.