Bodil A. Bluhm

Global Patterns and Predictions of Seafloor Biomass Using Random Forests

A comprehensive seafloor biomass and abundance database has been constructed from 24 oceanographic institutions worldwide within the Census of Marine Life (CoML) field projects. The machine-learning algorithm, Random Forests, was employed to model and predict seafloor standing stocks from surface primary production, water-column integrated and export particulate organic matter (POM), seafloor relief, and bottom water properties. The predictive models explain 63% to 88% of stock variance among the major size groups. Individual and composite maps of predicted global seafloor biomass and abundance are generated for bacteria, meiofauna, macrofauna, and megafauna (invertebrates and fishes). Patterns of benthic standing stocks were positive functions of surface primary production and delivery of the particulate organic carbon (POC) flux to the seafloor. At a regional scale, the census maps illustrate that integrated biomass is highest at the poles, on continental margins associated with coastal upwelling and with broad zones associated with equatorial divergence. Lowest values are consistently encountered on the central abyssal plains of major ocean basins The shift of biomass dominance groups with depth is shown to be affected by the decrease in average body size rather than abundance, presumably due to decrease in quantity and quality of food supply. This biomass census and associated maps are vital components of mechanistic deep-sea food web models and global carbon cycling, and as such provide fundamental information that can be incorporated into evidence-based management.

REGIONAL VARIABILITY IN FOOD AVAILABILITY FOR ARCTIC MARINE MAMMALS

This review provides an overview of prey preferences of seven core Arctic marine mammal species (AMM) and four non-core species on a pan-Arctic scale with regional examples. Arctic marine mammal species exploit prey resources close to the sea ice, in the water column, and at the sea floor, including lipid-rich pelagic and benthic crustaceans and pelagic and ice-associated schooling fishes such as capelin and Arctic cod. Prey preferred by individual species range from cephalopods and benthic bivalves to Greenland halibut. A few AMM are very prey-, habitat-, and/or depth-specific (e.g., walrus, polar bear), while others are rather opportunistic and, therefore, likely less vulnerable to change (e.g., beluga, bearded seal). In the second section, we review prey distribution patterns and current biomass hotspots in the three major physical realms (sea ice, water column, and seafloor), highlighting relations to environmental parameters such as advection patterns and the sea ice regime. The third part of the contribution presents examples of documented changes in AMM prey distribution and biomass and, subsequently, suggests three potential scenarios of large-scale biotic change, based on published observations and predictions of environmental change. These scenarios discuss (1) increased pelagic primary and, hence, secondary production, particularly in the central Arctic, during open-water conditions in the summer (based on surplus nutrients currently unutilized); (2) reduced benthic and pelagic biomass in coastal/shelf areas (due to increased river runoff and, hence, changed salinity and turbidity conditions); and (3) increased pelagic grazing and recycling in open-water conditions at the expense of the current tight benthic-pelagic coupling in part of the ice-covered shelf regions (due to increased pelagic consumption vs. vertical flux). Should those scenarios hold true, pelagic-feeding and generalist AMM might be advantaged, while the range for benthic shelf-feeding, ice-dependent AMM such as walrus would decrease. New pelagic feeding grounds may open up to AMM and subarctic marine mammal species in the High Arctic basins while nearshore waters might provide less abundant food in the future.

Food web structure in the high Arctic Canada Basin: evidence from δ13C and δ15N analysis

The food-web structure of the Arctic deep Canada Basin was investigated in summer 2002 using carbon and nitrogen stable isotope tracers. Overall food-web length of the range of organisms sampled occupied four trophic levels, based on 3.8‰ trophic level enrichment (δ15N range: 5.3–17.7‰). It was, thus, 0.5–1 trophic levels longer than food webs in both Arctic shelf and temperate deep-sea systems. The food sources, pelagic particulate organic matter (POM) (δ13C=−25.8‰, δ15N=5.3‰) and ice POM (δ13C=−26.9‰, δ15N=4.1‰), were not significantly different. Organisms of all habitats, ice-associated, pelagic and benthic, covered a large range of δ15N values. In general, ice-associated crustaceans (δ15N range 4.6–12.4‰, mean 6.9‰) and pelagic species (δ15N range 5.9–16.5, mean 11.5‰) were depleted relative to benthic invertebrates (δ15N range 4.6–17.7‰, mean 13.2‰). The predominantly herbivorous and predatory sympagic and pelagic species constitute a shorter food chain that is based on fresh material produced in the water column. Many benthic invertebrates were deposit feeders, relying on largely refractory material. However, sufficient fresh phytodetritus appeared to arrive at the seafloor to support some benthic suspension and surface deposit feeders on a low trophic level (e.g., crinoids, cumaceans). The enriched signatures of benthic deposit feeders and predators may be a consequence of low primary production in the high Arctic and the subsequent high degree of reworking of organic material.

Towards a pan-Arctic inventory of the species diversity of the macro- and megabenthic fauna of the Arctic shelf seas

Although knowledge of Arctic seas has increased tremendously in the past decade, benthic diversity was investigated at regional scales only, and no attempt had been made to examine it across the entire Arctic. We present a first pan-Arctic account of the species diversity of the macro- and megabenthic fauna of the Arctic marginal shelf seas. It is based on an analysis of 25 published and unpublished species-level data sets, together encompassing 14 of the 19 marine Arctic shelf ecoregions and comprising a total of 2,636 species, including 847 Arthropoda, 668 Annelida, 392 Mollusca, 228 Echinodermata, and 501 species of other phyla. For the four major phyla, we also analyze the differences in faunal composition and diversity among the ecoregions. Furthermore, we compute gross estimates of the expected species numbers of these phyla on a regional scale. Extrapolated to the entire fauna and study area, we arrive at the conservative estimate that 3,900–4,700 macro- and megabenthic species can be expected to occur on the Arctic shelves. These numbers are smaller than analogous estimates for the Antarctic shelf but the difference is on the order of about two and thus less pronounced than previously assumed. On a global scale, the Arctic shelves are characterized by intermediate macro- and megabenthic species numbers. Our preliminary pan-Arctic inventory provides an urgently needed assessment of current diversity patterns that can be used by future investigations for evaluating the effects of climate change and anthropogenic activities in the Arctic.

Freshwater and its role in the Arctic Marine System: Sources, disposition, storage, export, and physical and biogeochemical consequences in the Arctic and global oceans

The Arctic Ocean is a fundamental node in the global hydrological cycle and the oceans thermohaline circulation. We here assess the systems key functions and processes: (1) the delivery of fresh and low-salinity waters to the Arctic Ocean by river inflow, net precipitation, distillation during the freeze/thaw cycle, and Pacific Ocean inflows; (2) the disposition (e.g., sources, pathways, and storage) of freshwater components within the Arctic Ocean; and (3) the release and export of freshwater components into the bordering convective domains of the North Atlantic. We then examine physical, chemical, or biological processes which are influenced or constrained by the local quantities and geochemical qualities of freshwater; these include stratification and vertical mixing, ocean heat flux, nutrient supply, primary production, ocean acidification, and biogeochemical cycling. Internal to the Arctic the joint effects of sea ice decline and hydrological cycle intensification have strengthened coupling between the ocean and the atmosphere (e.g., wind and ice drift stresses, solar radiation, and heat and moisture exchange), the bordering drainage basins (e.g., river discharge, sediment transport, and erosion), and terrestrial ecosystems (e.g., Arctic greening, dissolved and particulate carbon loading, and altered phenology of biotic components). External to the Arctic freshwater export acts as both a constraint to and a necessary ingredient for deep convection in the bordering subarctic gyres and thus affects the global thermohaline circulation. Geochemical fingerprints attained within the Arctic Ocean are likewise exported into the neighboring subarctic systems and beyond. Finally, we discuss observed and modeled functions and changes in this system on seasonal, annual, and decadal time scales and discuss mechanisms that link the marine system to atmospheric, terrestrial, and cryospheric systems.

The autofluorescent age pigment lipofuscin: key to age, growth and productivity of the Antarctic amphipod Waldeckia obesa (Chevreux, 1905)

Peracarid crustaceans are among the most important taxa in terms of biodiversity and carbon-flow within the Weddell Sea benthos; however, very few data on their age, growth and productivity are available. This study uses the pigment lipofuscin as an age marker in the scavenging amphipod Waldeckia obesa (Chevreux, 1905) from the eastern Weddell Sea. Resin brain sections of 159 trap-caught specimens (1.2 to 7.7 mm coxal plate length L(cox) equal to 5 to 31 mm total length) were recorded digitally by confocal microscopy, and images were analysed. A modal progression analysis of the lipofuscin concentration-frequency distribution revealed five regularly spaced modes presumed to reflect consecutive annual age classes. Single females outside the range of mode V occurred, indicating maximum age of up to 8 years in females. No regular modes were obvious from the comparable length-frequency distribution of 386 individuals. Average yearly pigment accumulation was linear, and accumulation rates did not differ between sexes. The estimates of the growth parameters L(infinity) and k of the von Bertalanffy growth function were 7.47 mm L(cox) and 0.50 per year in females, respectively, and 6.92 mm L(cox) and 0.60 per year in males, respectively. Mortality, estimated from catch curves, amounted to 0.27 per year in females and 0.43 per year in males. P/B ratio, calculated from the mass specific growth rate method, was 0.38 per year for the pooled population (0.25 per year in females, 0.31 per year in males, 2.26 per year in juveniles). The results are discussed with regard to advantages and drawbacks of the methodology, and are compared with results from warmer water habitats.

Distribution, standing stock, growth, mortality and production of Strongylocentrotus pallidus (Echinodermata Echinoidea) in the northern Barents Sea

Abstract The regular sea urchin, Strongylocentrotus pallidus (G.O. Sars, 1871), is a widespread epibenthic species in high-Arctic waters. However, little is known about its distribution, standing stock, population dynamics and production. In the northern Barents Sea, S. pallidus was recorded on seabed still photographs at 10 out of 11 stations in water depths of 80–360 m. Mean abundances along photographic transects of 150–300 m length ranged between <0.1 and 14.7 ind. m−2 yielding a grand average of 3.6 ind. m−2. The small-scale distribution along the transects was patchy, with densities varying from nil to an overall maximum of 25.5 ind. m−2, and exhibited a significant relation to the number of stones present. Sea urchin test diameters, measured on scaled photographs, extended from 7 to 90 mm. Median values at single stations varied from 14 to 46 mm, showing a significant inverse relationship to water depth. Biomass, estimated by combining photographic abundances, size frequencies and a size-mass function established with trawled specimens, ranged between <0.1 and 3.0 g ash-free dry mass m−2, averaging about 1.0 g ash free dry mass m−2. An analysis of skeletal growth bands in genital plates was carried out with 143 trawled individuals ranging in test diameter (D) from 4 to 48 mm. Assuming these bands to represent annual growth marks, the ages of the specimens analysed ranged between 3 and 42 years. A von Bertalanffy function was fitted to size-at-age data to model individual growth pattern (D∞ = 102.3 mm, k = 0.011 year−1, t0 = 0.633 year). The annual mortality rate Z of the population in the northern Barents Sea was estimated from a size-converted catch curve to be 0.08 year−1. Applying the weight-specific growth rate method, the average P/B ratio and the mean annual production of this population were estimated as 0.07 year−1 and 0.076 g AFDM m−2 year−1, respectively. In conclusion, S. pallidus is characterized by slow growth, low mortality, high longevity and low productivity. Because of its relatively high biomass, it is considered to contribute significantly to total benthic standing stock and carbon flux in the study area.

Biodiversity and Biogeography of the Lower Trophic Taxa of the Pacific Arctic Region: Sensitivities to Climate Change

The lower trophic level taxa underpin the marine ecosystems of the Pacific Arctic Region (PAR). Recent field observations indicate that range shifts, and changes in the relative abundance of particular taxa have occurred within the last decade. Here we provide a region wide survey of the diversity and distribution of viruses, bacteria, archaea, auto- and heterotrophic protists, as well as metazoan zooplankton and benthic organisms in the PAR. Our aim is to provide a foundation for the assessment of the changes within the lower trophic level taxa of the PAR and to document such change when possible. Sensitivities to the effects of climate change are also discussed. Our vision is to enable data-based predictions regarding ecological succession in the PAR under current climate scenarios, and to deepen our understanding regarding what the future holds for higher trophic level organisms and the carbon cycle.

Editorial - Arctic Ocean Diversity: synthesis

The most fundamental attributes of marine ecosystems are their Communities associated species composition, along with their specific abundances and biomass. Processoriented understanding of rates and interactions within ecosystems hinges on this first-order descriptive framework. While we know the major species and understand their roles in many parts of the world ocean, the fragmented nature of discrete studies has not fostered synthetic approaches. The societal need for such basic information has increased in recent decades as major facets of the human footprint are altering marine communities around the globe (i.e. climate change, species invasions, fisheries effects, oil and gas exploration, tourism). To understand such change, biodiversity studies spanning species inventories to functional linkages between diversity and ecosystems are necessary. Within this context, as well as driven by simple human curiosity, the International Census of Marine Life (CoML) was launched in 2000 (Yarincik and O’Dor 2005). CoML grew to a global network of researchers in more than 80 nations engaged in a 10-year scientific initiative to assess and explain the diversity, distribution, and abundance of life in the oceans. CoML addressed the fundamental questions “What lived in the oceans in the past, what lives in the oceans now, and what will live in the oceans in the future” (McIntyre 2010). The Arctic component of CoML, the Arctic Ocean Diversity project (ArcOD), was launched in 2004 (Gradinger et al. 2010), with a sister project launched in the Antarctic shortly thereafter (Schiaparelli and Hopcroft 2011). Globally, gaps in biodiversity knowledge are greatest in areas where the logistics limit access. In this regard, the Arctic is understudied due the challenges of sampling in remote icecovered waters. There is increased urgency to fill these gaps because climate change effects are strongly expressed in the Arctic, as apparent from the rapid loss of its sea ice over the past decades. The ArcOD umbrella sought to inventory biodiversity in the Arctic sea ice, water column and sea floor (Fig. 1)—from the shallow shelves to the deep basins—using a three-level approach: compilation of existing data, taxonomic identification of existing samples, and new collections focusing on taxonomic and regional gaps. While ArcOD was initiated mainly by US-based and Russian scientists, over 100 scientists in a dozen nations have contributed to ArcODrelated efforts, including many conducted during the International Polar Year 2007–9. In October 2010, the Census reported ‘A decade of discovery’ across regions and realms at the Royal Society in London. This present special issue presents a core contribution of ArcOD’s synthesis and contains pieces originally presented in their preliminary form at the Arctic Frontiers meeting in January 2010 in Tromso, Norway in the ‘Marine Biodiversity under Change’ session. The articles in this and the subsequent issue (Hop et al. 2011) have a strong focus on biodiversity, on a species, community and/or habitat level. Articles in this issue are pan-Arctic in spatial coverage with international author teams from ten countries and more than 25 institutions contributing the required expertise and majority of the data. The contributions in this issue are arranged in taxonomic order and span from microbes to marine mammals. Most contain new synthetic numerical analyses, as well as reviews of current knowledge, contemporary perspectives, and several presently expected This article belongs to the special issue “Arctic Ocean Diversity Synthesis”

Do meio- and macrobenthic nematodes differ in community composition and body weight trends with depth?

Nematodes occur regularly in macrobenthic samples but are rarely identified from them and are thus considered exclusively a part of the meiobenthos. Our study compares the generic composition of nematode communities and their individual body weight trends with water depth in macrobenthic (>250/300 µm) samples from the deep Arctic (Canada Basin), Gulf of Mexico (GOM) and the Bermuda slope with meiobenthic samples (<45 µm) from GOM. The dry weight per individual (µg) of all macrobenthic nematodes combined showed an increasing trend with increasing water depth, while the dry weight per individual of the meiobenthic GOM nematodes showed a trend to decrease with increasing depth. Multivariate analyses showed that the macrobenthic nematode community in the GOM was more similar to the macrobenthic nematodes of the Canada Basin than to the GOM meiobenthic nematodes. In particular, the genera Enoploides, Crenopharynx, Micoletzkyia, Phanodermella were dominant in the macrobenthos and accounted for most of the difference. Relative abundance of non-selective deposit feeders (1B) significantly decreased with depth in macrobenthos but remained dominant in the meiobenthic community. The occurrence of a distinct assemblage of bigger nematodes of high dry weight per individual in the macrobenthos suggests the need to include nematodes in macrobenthic studies.