Introduction to the special issue on the biology of the circumpolar flaw lead (CFL) in the Amundsen Gulf of the Beaufort Sea (Arctic Ocean)

The sea-ice cover of the Arctic Ocean is decreasing in areal extent and volume more rapidly than model predictions of its response to continued warming of earth’s lower atmosphere. Reflecting this decrease is the spatial and temporal widening of the circumpolar flaw lead, a zone of open water and unconsolidated ice along the coastlines of the shallow seas that surround the deep Arctic Ocean basins. As part of the International Polar Year, the Circumpolar Flaw Lead System Study deployed the research icebreaker Amundsen from October 2007 to August 2008 in the flaw lead segment bordering Banks Island in the Canadian Beaufort Sea. With participation from Canada, the USA, Spain, Germany, China, Russia, the UK, Norway, Belgium, the Netherlands, France, and Sweden, the multi-disciplinary overwintering expedition logged 9,812 scientist-days in an effort to decipher key aspects of the ecosystem of the flaw lead. Highlights of the scientific achievements documented in this special issue on the biology of the circumpolar flaw lead system help to capture the depth, breadth, and novelty of this remarkable exploration.     

Pleistocene agglutinated foraminifera from the Lomonosov Ridge and Amundsen Basin, Arctic Basin. Initial report on piston cores 2177-5 (KAL) and 2176-3 (KAL)

Noncalcareous Pleistocene sediments of the Central Arctic Ocean contain sparse benthic foraminiferal assemblages consisting entirely of agglutinated taxa. Deep water agglutinated foraminifera are studied from two piston cores collected from the Lomonosov Ridge and Amundsen Basin [Cores PS 2177-5 (KAL) and 2176-3 (KAL)]. Core PS 2177-5 (KAL) contains an assemblage of 10 species, dominated by Cyclammina pusilla Brady, and is interpreted to reflect a bathyal environment with variable organic flux and nutrition levels. Core PS 2176-3 (KAL) in the Amundsen Basin yielded a very depauperate benthic foraminiferal assemblage. It is assumed that the environment was inhospitable for agglutinated foraminifera.     

Seasonal changes in substrate utilization patterns by bacterioplankton in the Amundsen Gulf (western Arctic)

Due to logistic difficulties, biological processes along the Arctic winter remain poorly known. In particular, carbon sources used by bacterioplankton have not been identified. A previous study in Franklin Bay suggested that polymers were one of the main substrates used by bacteria. During the Circumpolar Flaw Lead System Study, we analyzed metabolic capabilities of the heterotrophic bacterioplankton using Biolog MT2 MicroPlates^® amended with custom-selected substrates. Our purpose was to test whether the use of polymers was a peculiarity of Franklin Bay or a robust feature of the Arctic winter community. Seventeen stations were sampled in the Amundsen Gulf (western Arctic), a very dynamic area with heterogeneous ice conditions, from February to July 2008, at the surface (0–12 m), intermediate depths (20–70 m), and near the bottom (200 m). In winter, when nutrients and chlorophyll a concentrations were low, the number of substrates used was close to zero. In early spring, when the levels of chlorophyll a increased, so did the number of substrates used. This was followed by a 1-month period with no substrates used in April and May. Finally, the activity recovered toward the summer. Amino acids were occasionally used. However, the group of substrates most commonly used at all depths was carbohydrates, especially cellobiose, maltose, N-acetyl-d-glucosamine and glycogen. All these are either polymers or monomers derived from polymers. These results confirm that the heterotrophic bacterial assemblage relies heavily on polysaccharides for subsistence during the Arctic winter.     

Composition and distribution of mesozooplankton in the Nansen Basin,Arctic Ocean,during summer

Information about the large-scale zooplankton distribution in the Nansen Basin, that part of the Arctic Ocean most directly influenced by the inflow of Atlantic Water, is still scarce. During the Polarstern expedition ARK IV/3 in July/August 1987 zooplankton from the upper 500 m was sampled along two transects across the western Nansen Basin. Spatial variations in faunal composition, abundance, and biomass are described. Special emphasis is given to the distribution patterns and possible drifting routes of species advected into the Arctic Ocean through Fram Strait. Forty-five species and eight taxa not determined to species level were identified. Copepods clearly dominate in terms of species number, abundance, and biomass. Ostracods and chaetognaths comprise the most important groups of the non-copepod zooplankton. Maximum species numbers were recorded over the abyssal plain around 83°N. Total abundance and biomass decreased by more than 80% towards the northern part of the basin. Cluster analysis revealed two major faunal assemblages, broadly overlapping over the central Nansen Basin, and three principal types of spatial distribution. The well-defined zonation observed in the large-scale zooplankton distribution closely resembles the major hydrographic and topographic features. The opposite movement of the Polar Surface Water and the Atlantic Water in the southern Nansen Basin might cause pronounced deviations between the drifting routes of species with different water mass preferences advected through Fram Strait.     

Macro- and megabenthic communities in the high Arctic Canada Basin: initial findings

We investigated the macro- and megabenthos of the Canada Basin, one of the biologically least known areas of the Arctic deep sea. Macro-infauna was collected with 11 box cores at six stations ranging from 640 to 3,250 m water depth. Total abundance ranged from 3 to 265 individuals 0.04 m^–2 (75–6,625 individuals m^–2) and decreased with increasing depth. Biomass ranged from 0.04 to 228 mg wet weight 0.04 m^–2 (1–5,700 mg wet weight m^–2) and followed the same trend. Polychaetes, crustaceans and bivalves dominated the faunal densities, biomass and species numbers. At Northwind Ridge (800 and 1,800 m), apparently historic (~5,000 years BP) fish otolith deposits were abundant. The invertebrate epi-megafauna was qualitatively analyzed from 9.2 h of video and 853 still images collected at four remotely operated vehicle stations. The epifauna was dominated by polychaetes, crustaceans, echinoderms, cnidarians and fish, with most suspension-feeders occurring in the Northwind Ridge area. A total of 90 benthic invertebrate species/taxa from four biogeographic affinities were identified, including at least three new species of Isopoda. The evident low abundances and biomass are in agreement with findings from the Eurasian Arctic deep sea.     

Phylogeographic structure and mitochondrial DNA variation in true lemmings (Lemmus) from the Eurasian Arctic

The geographic pattern of mtDNA variation in lemmings from 13 localities throughout the Eurasian Arctic was studied by using eight restriction enzymes and sequencing of the cytochrome b region. These data are used to reveal the vicariant history of Lemmus , and to examine the effect of the last glaciation on mtDNA variation by comparing diversity in formerly glaciated areas to the diversity in non-glaciated areas. Phylogenetic congruence across different Arctic taxa and association between observed discontinuities, and probable Pleistocene barriers, suggest that glacial-interglacial periods were crucial in the vicariant history of Lemmus. Differences in amount of divergence (2.1–9.1%) across different historical barriers indicate chronologically separate vicariant events during the Quaternary. Populations from a formerly glaciated area are no less variable than those in the non-glaciated area. Regardless of glaciation history, no population structure and high haplotype diversity were found within geographic regions. The lack of population structure indicates that populations with high ancestral haplotype diversity shifted their distribution during the Holocene, and that lemmings tracked a changing environment during the Quaternary without reduction of effective population size.     

Macrobenthos composition,abundance and biomass in the Arctic Ocean along a transect between Svalbard and the Makarov Basin

Macrofauna has been sampled at 30 stations, at water depths of 1018–4478 m, along a transect extending between Northern Svalbard and the Makarov Basin, as a basis for understanding aspects of the benthic ecology of the Arctic Ocean. Species numbers, abundances and biomasses were extremely low, and generally varied between 0 to 11/0.02 m², 0 to 850 individuals/m², and 0 to 82.65 g/m², respectively. A total of 42 species was found. The Amphipod Jassa marmorata was the most common species. Both numbers and biomasses of suspension-feeding species increased towards the Lomonosov Ridge, probably due to lateral transport of organic material by deep currents along the ridge.     

Ice algal assemblages and vertical export of organic matter from sea ice in the Barents Sea and Nansen Basin (Arctic Ocean)

Algal communities and export of organic matter from sea ice were studied in the offshore marginal ice zone (MIZ) of the northern Barents Sea and Nansen Basin of the Arctic Ocean north of Svalbard by means of ice cores and short-term deployed sediment traps. The observations cover a total of ten stations within the drifting pack ice, visited over a period of 3 years during the period of ice melt in May and July. Maximum flux of particulate organic carbon and chlorophyll a from the ice at 1 m depth (1,537 mg C m⁻² per day and 20 mg Chl a m⁻² per day) exceeded the flux at 30 m by a factor of 2 during spring, a pattern that was reversed later in the season. Although diatoms dominated the ice-associated algal biomass, flagellates at times revealed similarly high biomass and typically dominated the exported algal carbon. Importance of flagellates to the vertical flux increased as melting progressed, whereas diatoms made the highest contribution during the early melting stage. High export of ice-derived organic matter and phytoplankton took place simultaneously in the offshore MIZ, likely as a consequence of ice drift dynamics and the mosaic structure of ice-covered and open water characteristic of this region.     

Structure and distribution of pelagic ostracods (Ostracoda: Myodocopa) in the Arctic Ocean

The study of the pelagic ostracod fauna of the Arctic Ocean based on materials collected by numerous Russian expeditions (1929–1993) and data from the literature showed the extreme poorness of the Arctic pelagic ostracod fauna, its mainly North Atlantic genesis and complete isolation from the Pacific fauna. Maximum ostracod abundance was observed in the epipelagic zone, and the greatest species diversity occurred in the relatively warm deep Atlantic layer throughout the year. To the north, east, and west of Franz Josef Land and Spitsbergen, the number of species and abundance indices of pelagic ostracods were decreased. In superficial water layers of the Central Arctic, maximum ostracod density and biomass were recorded in June and September. The best bioindicator of warm Atlantic water in the Arctic basin is Obtusoecia obtusata; and of cold polar water in the North Atlantic, Boroecia maxima.     

Zooplankton boom and ice amphipod bust below melting sea ice in the Amundsen Gulf, Arctic Canada

Early summer in the Arctic with extensive ice melt and break-up represents a dramatic change for sympagic–pelagic fauna below seasonal sea ice. As part of the International Polar Year-Circumpolar Flaw Lead system study (IPY-CFL), this investigation quantified zooplankton in the meltwater layer below landfast ice and remaining ice fauna below melting ice during June (2008) in Franklin Bay and Darnley Bay, Amundsen Gulf, Canada. The ice was in a state of advanced melt, with fully developed melt ponds. Intense melting resulted in a 0.3- to 0.5-m-thick meltwater layer below the ice, with a strong halocline to the Arctic water below. Zooplankton under the ice, in and below the meltwater layer, was sampled by SCUBA divers. Dense concentrations (max. 1,400 ind. m⁻³) of Calanus glacialis were associated with the meltwater layer, with dominant copepodid stages CIV and CV and high abundance of nauplii. Less abundant species included Pseudocalanus spp., Oithona similis and C. hyperboreus. The copepods were likely feeding on phytoplankton (0.5–2.3 mg Chl-a m⁻³) in the meltwater layer. Ice amphipods were present at low abundance (<10 ind. m⁻²) and wet biomass (<0.2 g m⁻²). Onisimus glacialis and Apherusa glacialis made up 64 and 51% of the total ice faunal abundance in Darnley Bay and Franklin Bay, respectively. During early summer, the autochthonous ice fauna becomes gradually replaced by allochthonous zooplankton, with an abundance boom near the meltwater layer. The ice amphipod bust occurs during late stages of melting and break-up, when their sympagic habitat is diminished then lost.     

Variability of protistan and bacterial communities in two Arctic fjords (Spitsbergen)

Krossfjorden and Kongsfjorden are Arctic fjords on the western side of Spitsbergen. These fjords share a common mouth to the open sea, and both are influenced by the input of sediment-rich glacial meltwater leading to decreased surface salinity, increased turbidity and decreased light penetration during summer. Earlier classical taxonomic studies had described the pelagic protistan composition of the Kongsfjorden during summer, revealing the dominance of flagellates of often unresolved taxonomic origin. Only little information existed on microbial eukaryote composition of the Krossfjorden as well as the bacterial composition of both fjords. The aim of the present study was to analyze and compare surface summertime protistan and bacterial communities in both fjords, using molecular approaches (16S and 18S rRNA DGGE, sequencing). Samples were collected three times a week from the central Kongsfjorden over a 1-month period. Additionally, 10 marine and 2 freshwater sites were sampled within a 1-week period in both Kongsfjorden and Krossfjorden. The central Kongsfjorden revealed a relatively stable protistan community over time with dinoflagellates, chlorophytes and small heterotrophs dominating. In contrast, the bacterial community varied over time and appeared to be correlated with the inflow of glacial meltwater. The Kongsfjorden and Krossfjorden were found to harbor distinctive bacterial and eukaryotic communities. We speculate that differences in glacial meltwater composition and fjord bathymetry affect the surface water properties and therefore the observed spatial variability in the community fingerprints.     

Environmental drivers of macroinvertebrate communities in high Arctic rivers (Svalbard)

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The Impact of the Latest Danian Event on Planktic Foraminiferal Faunas at ODP Site 1210 (Shatsky Rise,Pacific Ocean)

The marine ecosystem has been severely disturbed by several transient paleoenvironmental events (<200 kyr duration) during the early Paleogene, of which the Paleocene-Eocene Thermal Maximum (PETM, ~56 Ma) was the most prominent. Over the last decade a number of similar events of Paleocene and Eocene age have been discovered. However, relatively little attention has been paid to pre-PETM events, such as the “Latest Danian Event” ("LDE", ~62.18 Ma), specifically from an open ocean perspective. Here we present new foraminiferal isotope (δ¹³C, δ¹⁸O) and faunal data from Ocean Drilling Program (ODP) Site 1210 at Shatsky Rise (Pacific Ocean) in order to reconstruct the prevailing paleoceanographic conditions. The studied five-meter-thick succession covers ~900 kyr and includes the 200-kyr-lasting LDE. All groups surface dwelling, subsurface dwelling and benthic foraminifera show a negative δ¹³C excursion of >0.6‰, similar in magnitude to the one previously reported from neighboring Site 1209 for benthic foraminifera. δ¹⁸O-inferred warming by 1.6 to 2.8°C (0.4–0.7‰ δ¹⁸O measured on benthic and planktic foraminiferal tests) of the entire water column accompanies the negative δ¹³C excursion. A well stratified upper ocean directly before and during the LDE is proposed based on the stable isotope gradients between surface and subsurface dwellers. The gradient is less well developed, but still enhanced after the event. Isotope data are supplemented by comprehensive planktic foraminiferal faunal analyses revealing a dominance of Morozovella species together with Parasubbotina species. Subsurface-dwelling Parasubbotina shows high abundances during the LDE tracing changes in the strength of the isotope gradients and, thus, may indicate optimal living conditions within a well stratified surface ocean for this taxon. In addition, distinct faunal changes are reported like the disappearance of Praemurica species right at the base of the LDE and the continuous replacement of M. praeangulata with M. angulata across the LDE.     

High gene flow in polar cod (Boreogadus saida) from West-Svalbard and the Eurasian Basin

The current and projected environmental change of the Arctic Ocean contrasts sharply with the limited knowledge of its genetic biodiversity. Polar cod Boreogadus saida (Lepechin, 1774) is an abundant circumpolar marine fish and ecological key species. The central role of polar cod in the Arctic marine food web warrants a better understanding of its population structure and connectivity. In this study, the genetic population structure of 171 juveniles, collected from several fjords off West-Svalbard (Billefjorden, Hornsund and Kongsfjorden), the northern Sophia Basin and the Eurasian Basin of the Arctic Ocean, was analysed using nine DNA microsatellite loci. Genetic analyses indicated moderate to high genetic diversity, but absence of spatial population structure and isolation-by-distance, suggesting ongoing gene flow between the studied sampling regions. High levels of connectivity may be key for polar cod to maintain populations across wide spatial scales. The adaptive capacity of the species will be increasingly important to face challenges such as habitat fragmentation, ocean warming and changes in prey composition. In view of a limited understanding of the population dynamics and evolution of polar cod, a valuable next step to predict future developments should be an integrated biological evaluation, including population genomics, a life-history approach, and habitat and biophysical dispersal modelling.     

Benthic Oxygen Uptake in the Arctic Ocean Margins - A Case Study at the Deep-Sea Observatory HAUSGARTEN (Fram Strait)

The past decades have seen remarkable changes in the Arctic, a hotspot for climate change. Nevertheless, impacts of such changes on the biogeochemical cycles and Arctic marine ecosystems are still largely unknown. During cruises to the deep-sea observatory HAUSGARTEN in July 2007 and 2008, we investigated the biogeochemical recycling of organic matter in Arctic margin sediments by performing shipboard measurements of oxygen profiles, bacterial activities and biogenic sediment compounds (pigment, protein, organic carbon, and phospholipid contents). Additional in situ oxygen profiles were performed at two sites. This study aims at characterizing benthic mineralization activity along local bathymetric and latitudinal transects. The spatial coverage of this study is unique since it focuses on the transition from shelf to Deep Ocean, and from close to the ice edge to more open waters. Biogeochemical recycling across the continental margin showed a classical bathymetric pattern with overall low fluxes except for the deepest station located in the Molloy Hole (5500 m), a seafloor depression acting as an organic matter depot center. A gradient in benthic mineralization rates arises along the latitudinal transect with clearly higher values at the southern stations (average diffusive oxygen uptake of 0.49 ± 0.18 mmol O₂ m^-2 d^-1) compared to the northern sites (0.22 ± 0.09 mmol O₂ m^-2 d^-1). The benthic mineralization activity at the HAUSGARTEN observatory thus increases southward and appears to reflect the amount of organic matter reaching the seafloor rather than its lability. Although organic matter content and potential bacterial activity clearly follow this gradient, sediment pigments and phospholipids exhibit no increase with latitude whereas satellite images of surface ocean chlorophyll a indicate local seasonal patterns of primary production. Our results suggest that predicted increases in primary production in the Arctic Ocean could induce a larger export of more refractory organic matter due to the longer production season and the extension of the ice-free zone.     

Macrofauna communities of the Dogger Bank (central North Sea) in the late 1990s: spatial distribution,species composition and trophic structure

Macrofauna samples were taken in May 1996–1998 at 28 stations on the Dogger Bank (central North Sea) to describe species composition, spatial distribution and trophic structure of the macrofaunal communities. In general, five communities could be distinguished depending on the influence of different water masses, depth, type of sediment and food availability. Except for those species which feed in the sediment, macrofauna communities did not show any correlation to the measured sediment variables. In fact, macrofauna communities of the Dogger Bank were mainly structured by lateral food supply via frontal systems. This was indicated by the dominance of interface feeders foraging in the benthic boundary layer, which are able to switch from surface deposit to suspension feeding depending on flow conditions. The importance of sandlickers in the shallowest parts of the Dogger Bank hints at an important contribution of benthic primary production to the nutritional and energetic needs of the benthic community. Electronic Publication     

Phylogeographical lineages of Arctic grayling (Thymallus arcticus) in North America: divergence, origins and affinities with Eurasian Thymallus

The number and location of Arctic glacial refugia utilized by taxa during the Pleistocene are continuing uncertainties in Holarctic phylogeography. Arctic grayling (Thymallus arcticus) are widely distributed in freshwaters from the eastern side of Hudson Bay (Canada) west to central Asia. We studied mitochondrial DNA (mtDNA) and microsatellite DNA variation in North American T. arcticus to test for genetic signatures of survival in, and postglacial dispersal from, multiple glacial refugia, and to assess their evolutionary affinities with Eurasian Thymallus. In samples from 32 localities, we resolved 12 mtDNA haplotypes belonging to three assemblages that differed from each other in sequence by between 0.75 and 2.13%: a 'South Beringia' lineage found from western Alaska to northern British Columbia, Canada; a 'North Beringia' lineage found on the north slope of Alaska, the lower Mackenzie River, and to eastern Saskatchewan; and a 'Nahanni' lineage confined to the Nahanni River area of the upper Mackenzie River drainage. Sequence analysis of a portion of the control region indicated monophyly of all North American T. arcticus and their probable origin from eastern Siberian T. arcticus at least 3 Mya. Arctic grayling sampled from 25 localities displayed low allelic diversity and expected heterozygosity (H(E)) across five microsatellite loci (means of 2.1 alleles and 0.27 H(E), respectively) and there were declines in these measures of genetic diversity with distance eastward from the lower Yukon River Valley. Assemblages defined by mtDNA divergences were less apparent at microsatellite loci, but again the Nahanni lineage was the most distinctive. Analysis of molecular variance indicated that between 24% (microsatellite DNA) and 81% (mtDNA) of the variance was attributable to differences among South Beringia, North Beringia and Nahanni lineages. Our data suggest that extant North American Arctic grayling are more diverse phylogeographically than previously suspected and that they consist of at least three major lineages that originated in distinct Pleistocene glacial refugia. T. arcticus probably originated and dispersed from Eurasia to North America in the late to mid-Pliocene, but our data also suggest more recent (mid-late Pleistocene) interactions between lineages across Beringia.     

Mitochondrial DNA variation and the evolutionary history of chromosome races of collared lemmings (Dicrostonyx) in the Eurasian Arctic

Collared lemmings (Dicrostonyx) demonstrate extensive chromosome variation along their circumpolar distribution in the high Arctic. To reveal the history of this genus and the origin of chromosome races in the Palearctic, we studied the geographical pattern of mtDNA variation in lemmings from 13 localities by using eight tetranucleotide restriction enzymes. The main split in mtDNA phylogeny is at the Bering Strait and corresponds to the main chromosome division between the Beringian and the Eurasian groups of karyotypes. Nucleotide divergence estimate of 6.8% suggests that, despite the Bering Land Bridge, Palearctic and Nearctic forms have been separated since the mid-Pleistocene. Five distinct phylogenetic groups of mtDNA haplotypes, with average divergence of 1.5%, corresponding to geographical regions, were found along the Palearctic coast. Low nucleotide and haplotype diversity and a star-like phylogeny within phylogeographical groups of haplotypes suggest regional bottleneck events in the recent past, most probably due to warming events during the Holocene. There is congruence between phylogeographical pattern of mtDNA variation and geographical distribution of chromosome races; 69% of the total mtDNA variation is allocated among chromosome races. This congruence implies that historical events such as fragmentation and allopatric bottleneck events have been important for the origin of chromosome races. However, historical factors do not explain the fixed autosome fusions found to distinguish certain populations.