Abstract13 C tracer experiments were conducted at sites spanning the steep oxygen, organic matter, and biological community gradients across the Arabian Sea oxygen minimum zone, in order to quantify the role that benthic fauna play in the short-term processing of organic matter (OM) and to determine how this varies among different environments. Metazoan macrofauna and macrofauna-sized foraminiferans took up as much as 56 6 13 mg of added C m 22 (685 mg C m 22 added) over 2-5 d, and at some sites this uptake was similar in magnitude to bacterial uptake and/or total respiration. Bottom-water dissolved oxygen concentrations exerted a strong control over metazoan macrofaunal OM processing. At oxygen concentrations .7 mmol L 21 (0.16 ml L 21 ), metazoan macrofauna were able to take advantage of abundant OM and to dominate OM uptake, while OM processing at O 2 concentrations of 5.0 mmol L 21 (0.11 ml L 21 ) was dominated instead by (macrofaunal) foraminiferans. This led us to propose the hypothesis that oxygen controls the relative dominance of metazoan macrofauna and foraminifera in a threshold manner, with the threshold lying between 5 and 7 mmol L 21 (0.11 to 0.16 ml L 21 ). Large metazoan macrofaunal biomass and high natural concentrations of OM were also associated with rapid processing of fresh OM by the benthic community. Where they were present, the polychaete Linopherus sp. and the calcareous foraminiferan Uvigerina ex gr. semiornata, dominated the uptake of OM above and below, respectively, the proposed threshold concentrations of bottom-water oxygen.1 Present address: Danish Meteorological Institute, Lyngbyvej 100, 2100 Copenhagen, Denmark. AcknowledgmentsThe experimental work in this study was conducted aboard the RRS Charles Darwin. We thank Oli Peppe and Eric Breuer for running the lander and megacorer. We also thank two anonymous reviewers for their constructive feedback.
The Pakistan margin is characterized by a strong midwater oxygen minimum zone (OMZ) that intercepts the seabed at bathyal depths (150 to 1300 m). We investigated whether faunal abundance and diversity trends were similar among protists (foraminiferans and gromiids), metazoan macrofauna and megafauna along a transect (140-1850 m water depth) across the OMZ during the 2003 intermonsoon (March-May) and late/post monsoon (August-October) seasons. All groups exhibited some drop in abundance in the OMZ core (250-500 m water depth; O 2 : 0.10-0.13 ml l -1 = 4.46-5.80 μM l -1 ) but to differing degrees. Densities of foraminiferans >63 µm were slightly depressed at 300 m, peaked at 738 m, and were much lower at deeper stations. Foraminiferans >300 µm were the overwhelmingly dominant macrofaunal organisms in the OMZ core.Macrofaunal metazoans reached maximum densities at 140 m depth, with additional peaks at 850, 940 and 1850 m where foraminiferans were less abundant. The polychaete Linopherus sp. was responsible for a macrofaunal biomass peak at 950 m. Apart from large swimming animals (fish and natant decapods), metazoan megafauna were absent between 300 and 900 m (O 2 <0.14-0.15 ml l -1 = 6.25-6.69 μM l -1 ) but were represented by a huge, ophiuroid-dominated abundance peak at 1000 m (O 2 ~0.15-0.18 ml l -1 = 6.69-8.03μM l -1 ). Gromiid protists were confined largely to depths below 1150 m (O 2 >0.2 ml l -1 = 8.92 μM l -1 ). The progressively deeper abundance peaks for foraminiferans (>63 µm), Linopherus sp. and ophiuroids probably represent lower OMZ boundary edge effects and suggest a link between body size and tolerance of hypoxia. Macro-and megafaunal organisms collected between 800 and 1100 m were dominated by a succession of different taxa, indicating that the lower part of the OMZ is also a region of rapid faunal change. Species diversity was depressed in all groups in the OMZ core, but this was much more pronounced for macrofauna and megafauna than for foraminiferans. Oxygen levels strongly influenced the taxonomic composition of all faunal groups. Calcareous foraminiferans dominated the seasonally and permanently hypoxic sites (136-300 m); agglutinated foraminiferans were relatively more abundant at deeper stations where oxygen concentrations were >0.13 ml l -1 (= 5.80 μM l -1 ). Polychaetes were the main macrofaunal taxon within the OMZ; calcareous macrofauna and megafauna (decapods, 3 echinoderms) were rare or absent where oxygen levels were lowest. The rarity of larger animals between 300 and 700 m on the Pakistan margin, compared with the abundant macrofauna in the OMZ core off Oman, is the most notable contrast between the two sides of the Arabian Sea. This difference probably reflects the slightly higher oxygen levels and better food quality on the western side.
Permafrost peatlands contain globally important amounts of soil organic carbon, owing to cold conditions which suppress anaerobic decomposition. However, climate warming and permafrost thaw threaten the stability of this carbon store. The ultimate fate of permafrost peatlands and their carbon stores is unclear because of complex feedbacks between peat accumulation, hydrology and vegetation. Field monitoring campaigns only span the last few decades and therefore provide an incomplete picture of permafrost peatland response to recent rapid warming. Here we use a high-resolution palaeoecological approach to understand the longer-term response of peatlands in contrasting states of permafrost degradation to recent rapid warming. At all sites we identify a drying trend until the late-twentieth century; however, two sites subsequently experienced a rapid shift to wetter conditions as permafrost thawed in response to climatic warming, culminating in collapse of the peat domes. Commonalities between study sites lead us to propose a five-phase model for permafrost peatland response to climatic warming. This model suggests a shared ecohydrological trajectory towards a common end point: inundated Arctic fen. Although carbon accumulation is rapid in such sites, saturated soil conditions are likely to cause elevated methane emissions that have implications for climate-feedback mechanisms.
Sedimented hydrothermal vents, where hot, mineral-rich water flows through sediment, are poorly understood globally, both in their distribution and the ecology of individual vent fields. We explored macrofaunal community ecology at a sediment-hosted hydrothermal vent in the Southern Ocean. This is the first such study of these ecosystems outside of the Pacific and the furthest south (62 • S) of any vent system studied. Sedimentary fauna were sampled in four areas of the Bransfield Strait (Southern Ocean), with the aim of contrasting community structure between vent and non-vent sites. Macrofaunal assemblages were clearly distinct between vent and non-vent sites, and diversity, richness, and density declined toward maximum hydrothermal activity. This variation is in contrast to observations from similar systems in the Pacific and demonstrates the influence of factors other than chemosynthetic primary productivity in structuring infauna at deep-sea vent communities. Vent endemic fauna had limited abundance and were represented by a single siboglinid species at hydrothermally active areas, meaning that that the majority of local biota were those also found in other areas. Several taxa occupied all sampling stations but there were large differences in their relative abundances, suggesting communities were structured by niche variation rather than dispersal ability.
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