Anaerobic ammonium-oxidizing bacteria in marine environments: widespread occurrence but low diversity

Laboratory and field studies have indicated that anaerobic ammonium oxidation (anammox) is an important process in the marine nitrogen cycle. In this study 11 additional anoxic marine sediment and water column samples were studied to substantiate this claim. In a combined approach using the molecular methods, polymerase chain reaction (PCR), qualitative and quantitative fluorescence in situ hybridization (FISH), as well as (15)N stable isotope activity measurements, it was shown that anammox bacteria were present and active in all samples investigated. The anammox activity measured in the sediment samples ranged from 0.08 fmol cell(-1) day(-1) N(2) in the Golfo Dulce (Pacific Ocean, Costa Rica) sediment to 0.98 fmol cell(-1) day(-1) N(2) in the Gullmarsfjorden (North Sea, Sweden) sediment. The percentage of anammox cell of the total population (stained with DAPI) as assessed by quantitative FISH was highest in the Barents Sea (9% +/- 4%) and in most of the samples well over 2%. Fluorescence in situ hybridization and phylogenetic analysis of the PCR products derived from the marine samples indicated the exclusive presence of members of the Candidatus'Scalindua' genus. This study showed the ubiquitous presence of anammox bacteria in anoxic marine ecosystems, supporting previous observations on the importance of anammox for N cycling in marine environments.     

Insemination Capacity and Dispersal in Relation to Sex Allocation Decisions in Goniozus legneri (Hymenoptera: Bethylidae): Why Are There More Males in Larger Broods?

Models considering sex ratio optima under single foundress strict local mate competition predict that female bias will be reduced by stochasticity in sex allocation, developmental mortality of males and limited insemination capacity of males. In all three cases the number of males per brood is expected to increase with brood size. Sex ratio optima may also be less female biased when several mothers contribute offspring to local mating groups or if non‐local mating occurs between members of different broods; again more males are expected in larger broods. In the parasitoid wasp Goniozus legneri (Hymenoptera: Bethylidae), sex allocation has only a small stochastic component, developmental mortality is low and non‐siblings are unlikely to develop in the same brood. However, the number of males per brood increases with the size of the brood (produced by a single mother). We investigated the further possibilities of limited insemination capacity and non‐local mating using a naturalistic experimental protocol. We found that limited insemination capacity is an unlikely general explanation for the increase in number of males with brood size. All males and females dispersed from both mixed and single sex broods. Although most females in mixed sex broods mated prior to dispersal, these data suggest that non‐local mating is possible, for instance via male immigration to broods containing virgin females. This may influence sex ratio optima and account for the trend in male number.     

Macrobenthic species composition and diversity in the Godthaabsfjord system, SW Greenland

The southwest Greenland coast is made up of large and deep sill fjords. On the shelf, a number of shallow banks separated by deep troughs are located 20–50 km from the coast. We collected three 0.1-m² van Veen grabs at nine stations along a transect spanning from the inner Godthaabsfjord influenced by glaciers, across the shallow Fyllas Bank and out to the slope of the continental shelf at approximately 1,000 m depth. Along this transect, we explored patterns of macrobenthic diversity, species composition, abundance and biomass. The sampled stations were very different in terms of environmental variables, resulting in large differences in species composition primarily related to differences in depth, silt–clay fraction and chl a content of the sediment (BIO-ENV analysis). Habitat differences also reduced species spatial ranges and the majority of species were found at only one (49%) or two (20%) stations and, consequently, species turnover or beta diversity was high and correlated to differences in depth, silt–clay fraction and median sediment grain size. Species richness and diversity were lowest in sites exposed to sediment disturbance: near the glaciers in the inner fjord (physical disturbance by mineral sedimentation) and at selected stations on the shelf (bioturbation by burrowing sand eel). Alpha diversity and richness were only weakly correlated to environmental parameters, indicating that alpha richness and diversity are influenced by several factors or that relationships are non-linear as was found for species richness and silt–clay fraction.     

Abundance and energy requirements of eiders (Somateria spp.) suggest high predation pressure on macrobenthic fauna in a key wintering habitat in SW Greenland

The number of common eiders (Somateria mollissima borealis) in west Greenland declined dramatically during the twentieth century, supposedly because of human activities. However, their sensitivity to alternative drivers of variation, such as climate conditions, diseases or food availability, remains unstudied. In this study, we describe prey availability and assess the trophic coupling between eiders and their macrobenthic prey in a shallow inlet, Nipisat Sound; a key wintering habitat in the south-west Greenland Open Water Area. Macrobenthic species abundance and biomass were studied, and annual production was estimated by an empirical model, including environmental characteristics, fauna composition and individual biomass. In spring 2008, average macrozoobenthic abundance and biomass were 6,912 ind m⁻² and 28.4 g ash-free dry mass (AFDM) m⁻² (647 kJ m⁻²), respectively. Annual production was estimated at 13.9 g AFDM m⁻² year⁻¹ (317 kJ m⁻² year⁻¹). During the winters of 2008–2010, we monitored the number of common eiders (S. mollissima borealis) and king eiders (Somateria spectabilis) and observed a distinct peak in abundance during winter with up to 15.000 birds in Nipisat Sound. Based on physiological costs of different activities in combination with the observed behavioural pattern, we obtained an estimate of the energy required for eiders to balance their costs of living, which amounted to 58% of the estimated total annual production of macrobenthos in Nipisat Sound. This result suggests that eider predation affects macrobenthic species composition and biomass and demonstrates the potential importance of variations in prey availability for the population dynamics of eiders in Greenland.     

Underwater observations of foraging free-living Atlantic walruses (Odobenus rosmarus rosmarus) and estimates of their food consumption

Food consumption of Atlantic walruses (Odobenus rosmarus rosmarus L.) was quantified by combining underwater observations of feeding with satellite-telemetry data on movement and diving activity. The study was conducted between 31 July and 7 August 2001 in Young Sound (74°N-20°W) in Northeast Greenland. On ten occasions, divers were able to accompany foraging walruses to the sea floor and collect the shells of newly predated bivalves (Mya truncata, Hiatella arctica, Serripes groenlandicus) for determination of number of prey and biomass ingested per dive. Simultaneously, the activity of a 1,200-kg adult male walrus was studied by use of satellite-telemetry during an entire foraging cycle that included 74 h at sea followed by a 23-h rest on land. An average of 53.2 bivalves (SE=5.2, range: 34-89, n=10) were consumed per dive, corresponding to 149.0 g shell-free dry matter (SE=18.9, range: 62.4-253.1 g), or 2,576 kJ per dive (SE=325.2, range: 1,072-4,377 kJ). During the foraging trip, the walrus spent 57% of the time diving to depths of between 6 and 32 m, and it made a total of 412 dives that lasted between 5 and 7 min (i.e. typical foraging dives). If the entire feeding cycle is considered (97 h), the estimated daily gross energy intake was 214 kJ per kg body mass (95% CI: 153-275 kJ), corresponding to the ingestion of 57 kg (95% CI: 41-72 kg) wet weight bivalve biomass per day, or 4.7 (95% CI: 3.3-5.9%) of total walrus body mass. Due to ice cover, walrus access to the plentiful inshore bivalve banks in the area is restricted to the short summer period, where walruses rely on them for replenishing energy stores. It is hypothesised that the documented decrease in the extent and duration of Arctic sea ice may increase food availability for walruses in eastern Greenland in the future.     

Seasonal sea ice cover as principal driver of spatial and temporal variation in depth extension and annual production of kelp in Greenland

We studied the depth distribution and production of kelp along the Greenland coast spanning Arctic to sub‐Arctic conditions from 78 ºN to 64 ºN. This covers a wide range of sea ice conditions and water temperatures, with those presently realized in the south likely to move northwards in a warmer future. Kelp forests occurred along the entire latitudinal range, and their depth extension and production increased southwards presumably in response to longer annual ice‐free periods and higher water temperature. The depth limit of 10% kelp cover was 9–14 m at the northernmost sites (77–78 ºN) with only 94–133 ice‐free days per year, but extended to depths of 21–33 m further south (73 ºN–64 ºN) where >160 days per year were ice‐free, and annual production of Saccharina longicruris and S. latissima, measured as the size of the annual blade, ranged up to sevenfold among sites. The duration of the open‐water period, which integrates light and temperature conditions on an annual basis, was the best predictor (relative to summer water temperature) of kelp production along the latitude gradient, explaining up to 92% of the variation in depth extension and 80% of the variation in kelp production. In a decadal time series from a high Arctic site (74 ºN), inter‐annual variation in sea ice cover also explained a major part (up to 47%) of the variation in kelp production. Both spatial and temporal data sets thereby support the prediction that northern kelps will play a larger role in the coastal marine ecosystem in a warmer future as the length of the open‐water period increases. As kelps increase carbon‐flow and habitat diversity, an expansion of kelp forests may exert cascading effects on the coastal Arctic ecosystem.     

Effects of food concentration on clearance rate and energy budget of the Arctic bivalve Hiatella arctica (L) at subzero temperature

The influence of food concentration on clearance rate, respiration, assimilation, and excretion at −1.3 °C was studied on individuals of the bivalve Hiatella arctica (L.) from Young Sound, NE Greenland. Clearance rate, assimilation efficiency, respiration, and excretion rates were determined over a range of food concentrations using the microalga Rhodomonas baltica as food source. Physiological rates were generally low but responded significantly to increased food levels. Clearance rates and assimilation efficiency were reduced at increased food levels, whereas respiration and excretion increased. Assimilation efficiency was generally high, which may be an adaptation to the low food concentration during most of the year in NE Greenland. Low filtration rates limited ingestion rates and resulted in a low maximum assimilation of 3 J h⁻¹. Despite the low food intake, very low food concentrations were required for individual specimens to obtain a positive energy budget. Predicted growth based on rates of assimilation and respiration were compared to published estimates of annual growth in Young Sound. We estimate that 3 weeks of growth in the laboratory under optimal food conditions could match annual growth in situ. We interpret this as evidence that food limitation is the primary impediment to growth in this Arctic population.     

Marine‐terminating glaciers sustain high productivity in Greenland fjords

Accelerated mass loss from the Greenland ice sheet leads to glacier retreat and an increasing input of glacial meltwater to the fjords and coastal waters around Greenland. These high latitude ecosystems are highly productive and sustain important fisheries, yet it remains uncertain how they will respond to future changes in the Arctic cryosphere. Here we show that marine‐terminating glaciers play a crucial role in sustaining high productivity of the fjord ecosystems. Hydrographic and biogeochemical data from two fjord systems adjacent to the Greenland ice sheet, suggest that marine ecosystem productivity is very differently regulated in fjords influenced by either land‐terminating or marine‐terminating glaciers. Rising subsurface meltwater plumes originating from marine‐terminating glaciers entrain large volumes of ambient deep water to the surface. The resulting upwelling of nutrient‐rich deep water sustains a high phytoplankton productivity throughout summer in the fjord with marine‐terminating glaciers. In contrast, the fjord with only land‐terminating glaciers lack this upwelling mechanism, and is characterized by lower productivity. Data on commercial halibut landings support that coastal regions influenced by large marine‐terminating glaciers have substantially higher marine productivity. These results suggest that a switch from marine‐terminating to land‐terminating glaciers can substantially alter the productivity in the coastal zone around Greenland with potentially large ecological and socio‐economic implications.     

Biomineralization plasticity and environmental heterogeneity predict geographical resilience patterns of foundation species to future change

Although geographical patterns of species' sensitivity to environmental changes are defined by interacting multiple stressors, little is known about compensatory processes shaping regional differences in organismal vulnerability. Here, we examine large‐scale spatial variations in biomineralization under heterogeneous environmental gradients of temperature, salinity and food availability across a 30° latitudinal range (3,334 km), to test whether plasticity in calcareous shell production and composition, from juveniles to large adults, mediates geographical patterns of resilience to climate change in critical foundation species, the mussels Mytilus edulis and M. trossulus. We find shell calcification decreased towards high latitude, with mussels producing thinner shells with a higher organic content in polar than temperate regions. Salinity was the best predictor of within‐region differences in mussel shell deposition, mineral and organic composition. In polar, subpolar, and Baltic low‐salinity environments, mussels produced thin shells with a thicker external organic layer (periostracum), and an increased proportion of calcite (prismatic layer, as opposed to aragonite) and organic matrix, providing potentially higher resistance against dissolution in more corrosive waters. Conversely, in temperate, higher salinity regimes, thicker, more calcified shells with a higher aragonite (nacreous layer) proportion were deposited, which suggests enhanced protection under increased predation pressure. Interacting effects of salinity and food availability on mussel shell composition predict the deposition of a thicker periostracum and organic‐enriched prismatic layer under forecasted future environmental conditions, suggesting a capacity for increased protection of high‐latitude populations from ocean acidification. These findings support biomineralization plasticity as a potentially advantageous compensatory mechanism conferring Mytilus species a protective capacity for quantitative and qualitative trade‐offs in shell deposition as a response to regional alterations of abiotic and biotic conditions in future environments. Our work illustrates that compensatory mechanisms, driving plastic responses to the spatial structure of multiple stressors, can define geographical patterns of unanticipated species resilience to global environmental change.