An assessment of the production, distribution and fate of highly branched isoprenoid (HBI) biomarkers produced by sea ice and pelagic diatoms is necessary to interpret their detection and proportions in the northern Bering and Chukchi Seas. HBIs measured in surface sediments collected from 2012 to 2017 were used to determine the distribution and seasonality of the biomarkers relative to sea ice patterns. A northward gradient of increasing ice algae deposition was observed with localized occurrences of elevated IP 25 (sympagic HBI) concentrations from 68-70˚N and consistently strong sympagic signatures from 71-72.5˚N. A declining sympagic signature was observed from 2012 to 2017 in the northeast Chukchi Sea, coincident with declining sea ice concentrations. HBI fluxes were investigated on the northeast Chukchi shelf with a moored sediment trap deployed from August 2015 to July 2016. Fluxes of sea ice exclusive diatoms (Nitzschia frigida and Melosira arctica) and HBIproducing taxa (Pleurosigma, Haslea and Rhizosolenia spp.) were measured to confirm HBI sources and ice associations. IP 25 was detected year-round, increasing in March 2016 (10 ng m-2 d-1) and reaching a maximum in July 2016 (1331 ng m-2 d-1). Snowmelt triggered the release of sea ice algae into the water column in May 2016, while under-ice pelagic production contributed to the diatom export in June and July 2016. Sea ice diatom fluxes were strongly correlated with the IP 25 flux, however associations between pelagic diatoms and HBI fluxes were inconclusive. Bioturbation likely facilitates sustained burial of sympagic organic matter on the shelf despite the occurrence of pelagic diatom blooms. These results suggest that sympagic diatoms may sustain the food web through winter on the northeast Chukchi shelf. The reduced relative proportions of sympagic HBIs in the northern Bering Sea are likely driven by sea ice persistence in the region.

We studied ice algae utilization by benthic fauna from the northern Bering and Chukchi Seas using highly branched isoprenoid (HBI) biomarkers. We assessed whether various food acquisition strategies influence the observed HBI signatures. The proportion of phytoplankton to ice algae-sourced HBIs was determined through the H-Print approach that is presumed to reflect the percentage of sea ice organic carbon (iPOC) incorporated into tissues, relative to phytoplankton organic carbon. Cluster analysis separated 3 groups based on location and feeding strategy that were significantly influenced by annual sea ice persistence. Ice algae utilization was most significant in the northeast Chukchi Sea, where seasonal sea ice was present the longest. General feeding strategy was determined to be a significant factor in the degree of ice algae utilization. Predominant deposit feeders (both surface and subsurface) used more ice algae relative to suspension feeders. Organic carbon incorporated by predominant suspension feeders was primarily phytoplankton-based. The vast majority of all organisms sampled (~90%) incorporated a measurable quantity of iPOC. Sipunculids and brittle stars had the highest relative dependence on ice algae, while other taxa displayed plastic dietary responses, including the suspension/surface deposit feeder Macoma calcarea. This study indicates that ice algae are widely utilized in Pacific Arctic benthic food webs, but most benthic organisms displayed flexibility in consuming the available food sources. The elevated utilization of ice algae by deposit feeders may prove to be a disadvantage for these organisms if they cannot adapt to the ongoing decline of iPOC as seasonal sea ice declines.

The expected reduction of ice algae with declining sea ice may prove to be detrimental to the Pacific Arctic ecosystem. Benthic organisms that rely on sea ice organic carbon (iPOC) sustain benthic predators such as the Pacific walrus (Odobenus rosmarus divergens). The ability to track the trophic transfer of iPOC is critical to understanding its value in the food web, but prior methods have lacked the required source specificity. We analyzed the H-Print index, based on biomarkers of ice algae versus phytoplankton contributions to organic carbon in marine predators, in Pacific walrus livers collected in 2012, 2014 and 2016 from the Northern Bering Sea (NBS) and Chukchi Sea. We paired these measurements with stable nitrogen isotopes (δ15N) to estimate trophic position. We observed differences in the contribution of iPOC in Pacific walrus diet between regions, sexes, and age classes. Specifically, the contribution of iPOC to the diet of Pacific walruses was higher in the Chukchi Sea (52%) compared to the NBS (30%). This regional difference is consistent with longer annual sea ice persistence in the Chukchi Sea. Within the NBS, the contribution of iPOC to walrus spring diet was higher in females (~45%) compared to males (~30%) for each year (p < 0.001), likely due to specific foraging behavior of females to support energetic demands associated with pregnancy and lactation. Within the Chukchi Sea, the iPOC contribution was similar between males and females, yet higher in juveniles than in adults. Despite differences in the origin of organic carbon fueling the system (sea ice versus pelagic derived carbon), the trophic position of adult female Pacific walruses was similar between the NBS and Chukchi Sea (3.2 and 3.5, respectively), supporting similar diets (i.e. clams). Given the higher quality of organic carbon from ice algae, the retreat of seasonal sea ice in recent decades may create an additional vulnerability for female and juvenile Pacific walruses and should be considered in management of the species.

Sea ice primary production is considered a valuable energy source for Arctic marine food webs, yet the extent remains unclear through existing methods. Here we quantify ice algal carbon signatures using unique lipid biomarkers in over 2300 samples from 155 species including invertebrates, fish, seabirds, and marine mammals collected across the Arctic shelves. Ice algal carbon signatures were present within 96% of the organisms investigated, collected year-round from January to December, suggesting continuous utilization of this resource despite its lower proportion to pelagic production. These results emphasize the importance of benthic retention of ice algal carbon that is available to consumers year-round. Finally, we suggest that shifts in the phenology, distribution and biomass of sea ice primary production anticipated with declining seasonal sea ice will disrupt sympagic-pelagic-benthic coupling and consequently the structure and the functioning of the food web which is critical for Indigenous Peoples, commercial fisheries, and global biodiversity.