Bowhead whale (Balaena mysticetus) growth and feeding as estimated by δ13C techniques

Tissue ¹³C of bowhead whale, Balaena mysticetus, varies seasonally in response to geographic variations in the ¹³C of prey organisms consumed along the annual migratory route between the Bering, Chukchi and Beaufort Seas. Seasonal changes in body ¹³C of whales taken in 1986 from Alaskan waters provide a means of estimating energy intake from the different habitats in which the whales feed. Adult bowheads do not show significant seasonal shifts in the ¹³C of muscle and visceral fat. Their enrichment in ¹³C relative to subadult whales suggests that they acquire most of their food from fall and winter feeding or from unsampled parts of the summer range where zooplankton are enriched in ¹³C. Young individuals however, undergo marked seasonal shifts indicating that they feed heavily both in the summer and autumn or winter. Oscillations in ¹³C along the length of the baleen also provide a measure of age. Baleen growth rates decline as the whales age and provide a means of correcting for wear loss and allow aging of individuals less than 12 yr old. Although body length is a poor indicator of age in young bowhead whales, baleen length is closely correlated. Growth rates of bowheads after Year 1 are slow ( 0.4m yr^-1) and up to 20 yr are required to reach the assumed length of sexual maturity at 13 to 14m.     

Arctic marine mammal population status,sea ice habitat loss,and conservation recommendations for the 21st century

Arctic marine mammals (AMMs) are icons of climate change, largely because of their close association with sea ice. However, neither a circumpolar assessment of AMM status nor a standardized metric of sea ice habitat change is available. We summarized available data on abundance and trend for each AMM species and recognized subpopulation. We also examined species diversity, the extent of human use, and temporal trends in sea ice habitat for 12 regions of the Arctic by calculating the dates of spring sea ice retreat and fall sea ice advance from satellite data (1979–2013). Estimates of AMM abundance varied greatly in quality, and few studies were long enough for trend analysis. Of the AMM subpopulations, 78% (61 of 78) are legally harvested for subsistence purposes. Changes in sea ice phenology have been profound. In all regions except the Bering Sea, the duration of the summer (i.e., reduced ice) period increased by 5–10 weeks and by >20 weeks in the Barents Sea between 1979 and 2013. In light of generally poor data, the importance of human use, and forecasted environmental changes in the 21st century, we recommend the following for effective AMM conservation: maintain and improve comanagement by local, federal, and international partners; recognize spatial and temporal variability in AMM subpopulation response to climate change; implement monitoring programs with clear goals; mitigate cumulative impacts of increased human activity; and recognize the limits of current protected species legislation.     

Competition among penguins and cetaceans reveals trophic cascades in the western Ross Sea, Antarctica

An apparent trophic cascade that appears during summer in the western Ross Sea, Antarctica, explains why the Antarctic silverfish (Pleuragramma antarcticum) there becomes cannibalistic; its principal prey, crystal krill (Euphausia crystallorophias) becomes scarce; and the diatom community is minimally grazed compared to adjacent areas. The krill is the major grazer of diatoms. On the basis of fieldwork at Ross Island, we suggest that the cascade results from foraging by unusually numerous Adélie Penguins (Pygoscelis adeliae), minke whales (Balaenoptera bonaerensis), and fish-eating killer whales (Orcinus orca). These species and other top predators apparently deplete the krill and silverfish. In drawing our conclusions, we were aided by two "natural experiments." In one "experiment," large, grounded icebergs altered the seasonal pattern of change in regional sea-ice cover, but not the seasonal change in penguin diet and foraging behavior that was also detected during the pre-iceberg era. In the other "experiment," a short-term polynya (opening in the ice) brought penguins and whales together in a confined area, this time altering both penguin diet and foraging behavior. We conclude that the foraging of penguins and whales, and not a formerly hypothesized seasonal decrease in sea-ice cover, explains (1) the annual switch in the penguins' prey from krill to silverfish, (2) the subsequent lengthening of penguin foraging trips, and (3) a marked decline of cetaceans in the area later in the season. Reduction in the middle-trophic-level prey is expressed in the relaxed grazing pressure on phytoplankton.     

Time series analysis of elemental and isotopic data from biomineralized whale tissue

A temporal record of environmental conditions is often contained within accretionary biological tissue. These records can provide knowledge of the environmental conditions that existed at the time the tissue was formed. In this study, we look at trace element concentrations and isotopic ratios of carbon and nitrogen as contained in baleen from bowhead whales in the eastern and western Arctic Ocean. Time series techniques, including maximum likelihood method and likelihood ratio tests, are applied to analysis of data and inference about their mean structures.     

Vertical fine structure of the biomass and composition of algal communities in Arctic pack ice

The biomass and composition of algal communities in sea ice were studied during two summer expeditions to the central Arctic Ocean and the Greenland Sea. Based on algal pigment determination and cell counts, high biomass accumulations were found at the surface, in the interior and in the bottom layer of the ice floes. Pennate diatoms dominated in the bottom layer, while phototrophic flagellates and cysts of unknown origin were the most abundant taxa in the upper parts. The lowermost 20 to 40 cm contained between 4 and 62% of the entire algal biomass. Consequently, ice biological studies, which deal only with the bottom few centimetres of the ice floes, will underestimate algal biomass and production by factors of up to 25. Differences between the results of this study and published data from coastal locations point towards different biological regimes in Arctic sea ice. The algal biomass in coastal ice is about two orders of magnitude higher and composed mainly of diatoms, probably supported by nutrient influx from the water column. In the pack ice of the central Arctic, nutrient supply is probably reduced, and flagellates contribute substantially to total algal biomass. However, methodological problems might partially be responsible for the observed differences. Received: 12 June 1998 / Accepted: 11 December 1998     

Climate change threatens polar bear populations: a stochastic demographic analysis

The polar bear (Ursus maritimus) depends on sea ice for feeding, breeding, and movement. Significant reductions in Arctic sea ice are forecast to continue because of climate warming. We evaluated the impacts of climate change on polar bears in the southern Beaufort Sea by means of a demographic analysis, combining deterministic, stochastic, environment-dependent matrix population models with forecasts of future sea ice conditions from IPCC general circulation models (GCMs). The matrix population models classified individuals by age and breeding status; mothers and dependent cubs were treated as units. Parameter estimates were obtained from a capture-recapture study conducted from 2001 to 2006. Candidate statistical models allowed vital rates to vary with time and as functions of a sea ice covariate. Model averaging was used to produce the vital rate estimates, and a parametric bootstrap procedure was used to quantify model selection and parameter estimation uncertainty. Deterministic models projected population growth in years with more extensive ice coverage (2001-2003) and population decline in years with less ice coverage (2004-2005). LTRE (life table response experiment) analysis showed that the reduction in lambda in years with low sea ice was due primarily to reduced adult female survival, and secondarily to reduced breeding. A stochastic model with two environmental states, good and poor sea ice conditions, projected a declining stochastic growth rate, log lambdas, as the frequency of poor ice years increased. The observed frequency of poor ice years since 1979 would imply log lambdas approximately - 0.01, which agrees with available (albeit crude) observations of population size. The stochastic model was linked to a set of 10 GCMs compiled by the IPCC; the models were chosen for their ability to reproduce historical observations of sea ice and were forced with "business as usual" (A1B) greenhouse gas emissions. The resulting stochastic population projections showed drastic declines in the polar bear population by the end of the 21st century. These projections were instrumental in the decision to list the polar bear as a threatened species under the U.S. Endangered Species Act.     

Exploring the effects of reductions in krill biomass in the Southern Ocean on blue whales using a state-dependent foraging model

Many species of baleen whales were hunted to near extinction in the Southern Hemisphere. The recovery of these populations will be affected by the availability of krill, a major dietary component, in the Southern Ocean. We combine a novel energetics model for baleen whales with a state dependent foraging model to explore the impacts of an expanding krill fishery on baleen whales. We parameterize the model for blue whales, but with simple modifications it could be applied to most baleen whales. We predict that an expanding fishery will have a small but significant impact on the blue whale population through decreased birth rates. However, spreading the catch limit throughout the range of krill can reduce these effects. In addition, whales may be able to reduce these impacts through adaptive changes in foraging behavior. The relationship between krill abundance and blue whale foraging and reproductive success is nonlinear, such that larger reductions in krill biomass, potentially following a loss of sea ice due to climate change, could have a much larger negative impact on the recovery of blue whales.     

Light acclimation states of phytoplankton in the Southern Ocean, determined using photosynthetic pigment distribution

In high-latitude waters such as the Southern Ocean, the primary production of phytoplankton supports the ecosystem. To understand the photo-acclimation strategy of such phytoplankton within cold environments, the vertical distribution profile of photosynthetic pigments was analyzed in the Southern Ocean. Samples were taken along 110°E during the austral summer, and along 150°E and around the edge of the seasonal sea ice of the Antarctic Continent during the austral autumn. Pigment extraction methods were optimized for these samples. The standing crop of chlorophyll a was larger in the region along the edge of the seasonal sea ice than at sampling stations in open ocean areas. Chlorophyll concentration seemed to be dependent on the formation of thermo- and haloclines along the edge of the seasonal sea ice, but not in the open ocean where such clines are less pronounced. The marker pigments fucoxanthin and/or 19′-hexanoyloxyfucoxanthin were dominant at most sampling stations throughout the water column, while other marker pigments such as alloxanthin were quite low. This indicated that diatoms and/or haptophytes were the major phytoplankton in this area. Comparison of the relative ratio of fucoxanthin with that of 19′-hexanoyloxyfucoxanthin allowed some stations to be characterized as either diatom-dominant or haptophyte-dominant. The relative ratio of xanthophyll-cycle pigments (diadinoxanthin plus diatoxanthin) to chlorophyll a was high in surface waters and decreased gradually with depth. This suggests that near the ice edge during summer in the Southern Ocean, both diatoms and haptophytes acclimate to their light environments to protect their photosystems under high-light conditions.     

Structural confirmation of the sea ice biomarker IP25 found in Arctic marine sediments

IP₂₅ is a highly branched isoprenoid and an organic geochemical biomarker that is produced by some Arctic sea ice diatoms. IP₂₅ has previously been used in Arctic palaeo sea ice reconstruction studies and as a tracer for studying Arctic food webs. Here, the molecular structure of IP₂₅ has been confirmed by ¹H and ¹³C NMR spectroscopy following large-scale extraction from marine sediments obtained from the Canadian Arctic and purification using a combination of open-column and HPLC chromatographic methods. The structure of IP₂₅ was consistent between the three different sampling locations and was identical to that found previously for this biomarker following synthesis from a closely related highly branched isoprenoid diene. Since this study represents the first structural characterisation of IP₂₅ in sediments, future analysis of sedimentary IP₂₅ for palaeo Arctic sea ice reconstructions can be carried out with much greater confidence.     

Abundance and variability of microorganisms and transparent exopolymer particles across the ice–water interface of melting first-year sea ice in the Laptev Sea (Arctic)

The distribution and abundance of transparent exopolymer particles (TEP) was determined in and below pack ice of the Laptev Sea from July to September 1995. Samples were collected from the lowermost 10 cm of ice floes and at 10 cm below the ice–water interface. Abundance of bacteria, protists and TEP was determined, and the sea ice–water boundary layer was characterized using temperature, salinity and molecular viscous shear stress. TEP, with a distinct size distribution signal, were found in highest concentrations inside the sea ice, ranging from not detectable to 16 cm² l⁻¹ (median: 2.9 cm² l⁻¹). In the water, concentrations were one order of magnitude lower, ranged from below detection to 2.7 cm² l⁻¹ (median: 0.2 cm² l⁻¹) and decreased after the middle of August, whereas abundances of autotrophic flagellates (AF), diatoms, heterotrophic flagellates (HF) and ciliates increased. The abundance of TEP decreased with its size in all samples following a power law relationship. The relation of TEP to the microbial community differed between the sea ice and water, being positively correlated with bacteria and diatoms in the ice and negatively correlated with HF in the sea water. The presence of a pycnocline significantly influenced the abundance of organisms, diatom composition and TEP concentrations. Pennate diatoms dominated by Nitzschia frigida were most abundant inside the ice. Though bacteria have the potential to produce exopolymeric substances (EPS), the results of this study indicate that the majority of TEP at the ice–water interface in first-year Arctic summer pack ice are produced by diatoms. Received: 19 August 1999 / Accepted: 4 July 2000     

Carbon-isotope ratio gradients in western arctic zooplankton

Zooplankton from 87 stations in the Bering, Chukchi and Beaufort Seas sampled in 1985, 1986 and 1987 showed a geographic gradient in stable carbon-isotope ratios (¹³C). The zooplankton most depleted in ¹³C were found in the central and eastern Beaufort Sea and those most enriched were from the Bering and Chukchi Seas. Average ¹³C values ranged from-20.9 to-26.7 for copepods and from-19.4 to-25.1 for euphausiids. Euphausiids show a minimum of 1.0 enrichment relative to copepods throughout the study area. Relative biomasses of the major zooplankton taxa varied significantly across the Alaskan Beaufort Sea in October 1986, with euphausiids dominating in the west and copepods in the east. These differences in taxonomic composition affected the weighted ¹³C values for total zooplankton and may produce an even more pronounced geographic gradient in zooplankton ¹³C than that found within a single taxon. The bowhead whale Balaena mysticetus migrates between wintering areas in the Bering Sea and summering areas in the Beaufort Sea and feeds over this geographic range. The zooplankton ¹³C gradient is the probable source of ¹³C oscillations found along the baleen plates of this planktivore.     

Sea-bird aggregation at a deep North Pacific seamount

During June 1991, we studied sea birds at a mid-ocean seamount (Fieberling Guyot) in the eastern North Pacific Ocean. Avifaunal composition changed from small Procellariiformes [a storm-petrel; Oceanodroma leucorhoa (Vieillot)] away from the seamount to an assemblage dominated by larger tubenoses [mostly black-footed albatross Diomedea nigripes Audubon and Cook's petrel Pterodroma cookii (Gray)]. Compared to adjacent waters, sea-bird density and biomass within a 30-km radius centered on the seamount summit were 2.4 and 8 times higher, respectively. Individual sea-bird taxa were 2 to 40 times more abundant at the seamount relative to values reported previously from large-scale surveys of deep-ocean regions in the central North Pacific. In September 1991 we studied potential prey of sea birds in the upper water column using a neuston net and multiple opening-closing net system (MOCNESS) tows. Most potential prey types in the neuston exhibited no significant enhancement over the seamount. MOCNESS samples at 10 m depth, however, showed several prey types to be more abundant over the seamount, and the dominant size class of fish was slightly larger. We attribute the sea-bird aggregation observed at this seamount to changes in the abundance and/or behavior of pelagic organisms in the deep scattering layer (not adequately sampled in this study), perhaps augmented by migrations of seamount residents into the surface layers. Processes on and in the vicinity of seamounts may provide spatially-predictable prey to wide-ranging aerial sea birds foraging in this relatively austere environment.     

Trophic structure of coastal Antarctic food webs associated with changes in sea ice and food supply

Predicting the dynamics of ecosystems requires an understanding of how trophic interactions respond to environmental change. In Antarctic marine ecosystems, food web dynamics are inextricably linked to sea ice conditions that affect the nature and magnitude of primary food sources available to higher trophic levels. Recent attention on the changing sea ice conditions in polar seas highlights the need to better understand how marine food webs respond to changes in such broad-scale environmental drivers. This study investigated the importance of sea ice and advected primary food sources to the structure of benthic food webs in coastal Antarctica. We compared the isotopic composition of several seafloor taxa (including primary producers and invertebrates with a variety of feeding modes) that are widely distributed in the Antarctic. We assessed shifts in the trophic role of numerically dominant benthic omnivores at five coastal Ross Sea locations. These locations vary in primary productivity and food availability, due to their different levels of sea ice cover, and proximity to polynyas and advected primary production. The delta15N signatures and isotope mixing model results for the bivalves Laternula elliptica and Adamussium colbecki and the urchin Sterechinus neumeyeri indicate a shift from consumption of a higher proportion of detritus at locations with more permanent sea ice in the south to more freshly produced algal material associated with proximity to ice-free water in the north and east. The detrital pathways utilized by many benthic species may act to dampen the impacts of large seasonal fluctuations in the availability of primary production. The limiting relationship between sea ice distribution and in situ primary productivity emphasizes the role of connectivity and spatial subsidies of organic matter in fueling the food web. Our results begin to provide a basis for predicting how benthic ecosystems will respond to changes in sea ice persistence and extent along environmental gradients in the high Antarctic.     

Schadstoff-Ferntransport in die Arktis

The Arctic is still considered as one of the few unpolluted regions in the world. This is true if one compares the Arctic region with middle latitude regions which are influenced more by human activities. However, the Arctic region is not isolated from all human impacts. In recent years, high concentrations of persistent pollutants (organic chemicals, metals) were detected in top predators of the Arctic food chain and indigenous peoples from the Canadian and Greenland Arctic, although no local contamination sources are known. The comprehensive, scientific investigation of the past 20 years confirmed that the combination of atmospheric and waterborne long-range transport is the major source of the high concentrations of persistent organic pollutants (POPs) in the pristine Arctic environment. However, also pelagic marine organisms (e.g. Atlantic cod, marine mammals) can transport large amounts of persistent pollutants in their lipids and introduce contaminants into the Arctic food web. Thus, the pollutants are transported into the Arctic and subsequently accumulated through the short and unbranched Arctic food web of the top predators. The most accepted theory nowadays describes the long-range transport of persistent pollutants as a combination of atmospheric and sea current transport, or as a ‘global distillation’ process. Depending on such physical properties of the substances as vapour pressure and the ambient tempeature, persistent (semivolatile) contaminants are transported over different distances prior to deposition (sea surface, sediment, soil). After the deposition, however, and depending on the weather conditions and surrounding temperature, persistent pollutants will be re-evaporated into the atmosphere and undergo further atmospheric transport to the Arctic region. This process is also called the ‘grasshopper effect’. The global transport of persistent pollutants into Arctic regions can be described as a repeatedly occurring combination of atmospheric and waterborne transport in which the main transport vehicle depends on the physical properties of the transported compound. The role of characteristic meteorological conditions in the respective climate zones through which the contaminant is transported must not be underestimated. Strong seasonal differences in temperature and precipitation rule the global weather situation. Therefore, seasonal pattern differences occur for the distribution of some persistent pollutants in the Arctic environment depending on average temperature, main wind and sea current directions, humidity and day time light conditions (causing photochemical degradation). The consumption of traditionally hunted marine mammals (seals, whales) was identified as one of the main reasons for high contamination burdens in the Canadian and Greenland Inuit populations. Consequences and counter measures against high contamination loads in the Arctic human populations and ecosystems are currently under discussion. However, no comprehensive measures concerning restrictions of hunting traditions are taken by the respective governments to date due to the primary social consequences which are to be expected. The advising experts argue that such a restriction would destroy the original social structures of the Inuit populations. Therefore, the drawbacks of such a hunting restriction would weigh heavier than the expected positive effects of the reduction of contaminant burdens ‘Arctic dilemma’.     

A ciliate red tide at Barrow,Alaska

In September 1968 the first occurrence of extensive red water in the Arctic Ocean in the vicinity of Point Barrow, Alaska, was recorded. The organism causing this water discoloration was a fairly large (100 to 150 ) ciliated protozoan, with chlorophyll-containing endosymbionts. This ciliate, which is not identical to the commonly reported Mesodinium or Cyclotrichium species, is described in this paper, but cannot be identified with any organism described in the literature. Concentrations of dissolved inorganic nutrients and trace metals were determined on water samples obtained from the red water and also in the clear water adjacent to it. These data are discussed relative to the hydrographic conditions in the Chuckchi Sea around Point Barrow. A direct species' analysis by microscopic methods indicated that the plankton were quite similar in all the water samples except for the aforementioned red ciliate, which accounted for over 90% of the total cellular organic carbon in the samples from the red water. Chemical analyses indicated that the red tide ciliate contained approximately 51% protein, 33% lipid, 8% carbohydrate, 0.2% chlorophyll a, and 1.2% DNA.     

Spatial trends in abundance of long-finned pilot whales, white-beaked dolphins and harbour porpoises in West Greenland

Data from an aerial line transect survey conducted off West Greenland during August–September 2007 were used to estimate the abundance of long-finned pilot whales (Globicephala melas), white-beaked dolphins (Lagenorhynchus albirostris) and harbour porpoises (Phocoena phocoena). The abundance of each species was estimated using mark-recapture distance sampling techniques to correct for perception bias, and correction factors for time spent at the surface were applied. The fully corrected abundance estimates were 8,133 long-finned pilot whales, 11,984 white-beaked dolphins and 33,271 harbour porpoises. Based on density surface modelling methods, a count model with a generalised additive model formulation was used to relate abundance to spatial variables. Response curves indicated that the preferred habitats were deep offshore areas in Midwest Greenland for pilot whales, deep water over steep seabed slopes in South Greenland for white-beaked dolphins and relatively shallow inshore waters in Midwest–South Greenland for harbour porpoises. The abundance estimates and spatial trends for the three species are the first obtained from Greenland.     

Contrasting foraging tactics by northern gannets (<Emphasis Type="Italic">Sula bassana</Emphasis>) breeding in different oceanographic domains with different prey fields

In order to forage and to provision offspring effectively, seabirds negotiate a complex of behavioural, energetic, environmental and social constraints. In first tests of GPS loggers with seabirds in North America, we investigated the foraging tactics of free-ranging northern gannets (Sula bassana) at a large and a medium-sized colony that differed in oceanography, coastal position and prey fields. Gannets at Low Arctic colony (Funk Island) 50 km off the northeast coast of Newfoundland, Canada provisioned chicks almost entirely with small forage fish (capelin Mallotus villosus, 89%), while at boreal colony (Bonaventure Island) 3 km from shore in the Gulf of St. Lawrence, Quebec, Canada, large pelagic fish dominated parental prey loads (Atlantic mackerel Scomber scombrus 50%, Atlantic herring Clupea harengus 33%). Mean foraging range and the total distance travelled per foraging trip were significantly greater at the larger inshore colony (Bonaventure) than at the smaller offshore colony (Funk Island; 138 and 452 km vs. 64 and 196 km, respectively). Gannets from Funk Island consistently travelled inshore to forage on reproductive capelin shoals near the coast, whereas foraging flights of birds from Bonaventure were much more variable in direction and destination. Birds from the Low Arctic colony foraged in colder sea surface water than did birds from the boreal colony, and dive characteristics differed between colonies, which is concordent with the difference in prey base. Differences between the colonies reflect oceanographic and colony-size influences on prey fields that shape individual foraging tactics and in turn generate higher level colony-specific foraging “strategies”.     

A new source of dimethylsulfide (DMS) for the arctic atmosphere: ice diatoms

We report the first evidence that pennate diatoms growing within the bottom layer of first-year ice in the Arctic produce significant amounts of particulate dimethylsulfoniopropionate (DMSP_p) and dissolved DMSP+DMS. In 1992 in Resolute Passage, a tributary of Barrow Strait, DMSP_p concentrations within the bottom layer of ice reached 1055 mg S m^-3 at the end of the vernal bloom, a value one order of magnitude higher than the maximum value reported in antarctic ice. Bottom-ice concentrations in DMSP_p and DMSP_d+DMS were significantly correlated with the abundance of the dominant pennate diatom Nitzschia frigida. Intracellular concentration in DMSP of ice algae was very low (0.001 pg cell^-1) at the end of April when algae were light-limited and reached 1.17 pg cell^-1 in mid-May following an increase in light and algal growth. We calculate that the rapid release of the dissolved DMSP+DMS from the ice into surface waters following the ice break-up will generate a sea-to-air DMS flux of 0.7 mg S m^-2 d^-1, a pulse ten times higher than the mean arctic summer flux. We estimate that this 1-d pulse represents up to 5% of the annual DMS emission in the Arctic.     

Killer whale predation on marine mammals at Punta Norte,Argentina; food sharing,provisioning and foraging strategy

Summary The social dynamics of killer whales (Orcinus orca) that hunt marine mammals are apparently highly flexible, though strong individual associations do exist. The killer whales at Punta Norte offer an unusually detailed view of association patterns and foraging behaviour, and suggest a pattern of behaviour that optimizes hunting efficiency with exception only to strong associations between some individuals and the provisioning and training of offspring. The main points from this paper are as follows: First, hunting effort was concentrated where the capture rate was greatest. All pods selectively attacked the prey type for which they had the highest capture rate. The amount of southern sea lion prey captured was approximately equal to the estimated minimum energetic requirement for killer whales based on weight. Secondly, one whale in each pod did the majority of the hunting, and then provisioned the others in the pod. It was clear on numerous occasions that food was shared. A review of reported incidences of killer wales taking marine mammal prey suggests that it is common for a subset of the individuals in a pod to hunt. These results are discussed in the context of the evolution of foraging behaviour.Offprint requests to: A.R. Hoelzel at the first address