Deep-sea Benthic Ecosystems Research Articles

Climatic forcing of quaternary deep-sea benthic communities in the North Pacific Ocean

There is growing evidence that changes in deep-sea benthic ecosystems are modulated by climate changes, but most evidence to date comes from the North Atlantic Ocean. Here we analyze new ostracod and published foraminiferal records for the last 250,000 years on Shatsky Rise in the North Pacific Ocean. Using linear models, we evaluate statistically the ability of environmental drivers (temperature, productivity, and seasonality of productivity) to predict changes in faunal diversity, abundance, and composition. These microfossil data show glacial-interglacial shifts in overall abundances and species diversities that are low during glacial intervals and high during interglacials. These patterns replicate those previously documented in the North Atlantic Ocean, suggesting that the climatic forcing of the deep-sea ecosystem is widespread, and possibly global in nature. However, these results also reveal differences with prior studies that probably reflect the isolated nature of Shatsky Rise as a remote oceanic plateau. Ostracod assemblages on Shatsky Rise are highly endemic but of low diversity, consistent with the limited dispersal potential of these animals. Benthic foraminifera, by contrast, have much greater dispersal ability and their assemblages at Shatsky Rise show diversities typical for deep-sea faunas in other regions.Statistical analyses also reveal ostracod-foraminferal differences in relationships between environmental drivers and biotic change. Rarefied diversity is best explained as a hump-shaped function of surface productivity in ostracods, but as having a weak and positive relationship with temperature in foraminifera. Abundance shows a positive relationship with both productivity and seasonality of productivity in foraminifera, and a hump-shaped relationship with productivity in ostracods. Finally, species composition in ostracods is influenced by both temperature and productivity, but only a temperature effect is evident in foraminifera. Though complex in detail, the global-scale link between deep-sea ecosystems and Quaternary climate changes underscores the importance of the interaction between the physical and biological components of paleoceanographical research for better understanding the history of the biosphere.

Reliability of CARD-FISH Procedure for Enumeration of Archaea in Deep-Sea Surficial Sediments

The enumeration of Archaea in deep-sea sediment samples is still limited, although different methodological procedures have been applied. Among these, catalysed reporter deposition-fluorescence in situ hybridisation (CARD-FISH) technique is a promising tool for estimation of archaeal abundance in deep-sea sediment samples. Comparing different permeabilisation treatments, the best results obtained both on archaeal pure cultures and on natural assemblages were with hydrochloric acid (0.1M) and proteinase K (0.004U/ml) treatments. The application of CARD-FISH on deep-sea sediments revealed that Archaea reach up to 41% of total prokaryotic cells. Specific probes for planktonic Archaea showed that marine Crenarchaea dominated archaeal seafloor communities. No clear bathymetric trends were observed for archaeal abundances and the morphology of continental margin (slope vs. canyon) seems not to have a direct influence on archaeal relative abundances. The site-specific sediment habitat-both abiotic environmental setting and sedimentary organic matter quality-explain up to 65% of variance of archaeal, crenarchaeal and euryarchaeal relative abundance, suggesting a wide ecophysiological adaptation to deep-sea benthic ecosystems. The findings demonstrate that Archaea are an important component of benthic microbial assemblages so far neglected, and hence they lay the groundwork for more focused research on their ecological importance in the functioning of deep-sea benthic ecosystems.

Climatic forcing of eastern Mediterranean deep-water formation and benthic ecosystems during the past 22 000 years

Lateglacial and Holocene faunal and stable-isotope records from benthic foraminifers in the eastern Mediterranean Sea (EMS) suggest a high spatiotemporal variability of deep-water oxygenation and biogeochemical processes at the sea floor during that time. Changes in the oxygenation and food availability of the deep-sea ecosystems are closely linked to the hydrology of the EMS borderlands; they reflect orbital and suborbital climate variations of the high northern latitudes and the African monsoon system. During the last glacial maximum, cool surface waters and high evaporation resulted in maximum convection and oxic deep-waters in all sub-basins. Strong wind-induced mixing fostered surface-water production with seasonal phytodetritus fluxes. During the glacial termination and the Holocene, oxygenation and food availability of deep-sea benthic ecosystems were characterized by a pronounced regional differentiation. Local deep-water formation and trophic conditions were particularly variable in the northern Aegean Sea as a response to changes in riverine runoff and Black Sea outflow. During the interval of sapropel S1 formation in the early Holocene, average oxygen levels decreased exponentially with increasing water depth, suggesting a basin-wide shallowing of vertical convection superimposed by local signals. In the northernmost Aegean Sea, deep-water ventilation persisted during the early period of S1 formation, owing to temperature-driven local convection and the absence of low-salinity Black Sea outflow. At the same time, severe temporary anoxia occurred in the eastern Levantine basin at water depths as shallow as 900 m. This area was likely influenced by enhanced nutrient input of the Nile river that resulted in high organic matter fluxes and related high oxygen-consumption rates in the water column. In the southern Aegean and Levantine Seas, we observe a gradual increase in deep-water residence times, preceding S1 formation by approximately 1–1.5 kyr. Once oxygen levels fell below a critical threshold, the benthic ecosystems collapsed almost synchronously with the onset of S1 deposition. The recovery of benthic ecosystems during the terminal phase of S1 formation is controlled by subsequently deeper convection and re-ventilation over a period of approximately 1500 years. After the re-ventilation of the various sub-basins had been completed during the middle and late Holocene, deep-water renewal was more or less similar to recent rates. During that time, deep-sea ecosystem variability was driven by short-term changes in food quantity and quality as well as in seasonality, all of which are linked to millennial-scale changes in riverine runoff and associated nutrient input.

High densities and depth-associated changes of epibenthic megafauna along the Aleutian margin from 2000–4200 m

The Aleutian margin is a dynamic environment underlying a productive coastal ocean and subject to frequent tectonic disturbance. In July 2004, we used over 500 individual bottom images from towed camera transects to investigate patterns of epibenthic megafaunal density and community composition on the contiguous Aleutian margin (53°N 163°W) at depths of 2000 m, 3200 m and 4200 m. We also examined the influence of vertical isolation on the megafaunal assemblage across a topographic rise at 3200 m, located 30 km from the main margin and elevated 800 m above the surrounding seafloor. In comparison to previous reports from bathyal and abyssal depths, megafaunal densities along the Aleutian margin were remarkably high, averaging 5.38±0.43 (mean±1 standard error), 0.32±0.02 to 0.43±0.03 and 0.27±0.01 individuals m−2 at 2000 m, 3200 m and 4200 m, respectively. Diversity at 2000 m was elevated by 15–30% over the deeper sites (3200–4200 m) depending on the metric, while evenness was depressed by ~10%. Levels of richness and evenness were similar among the three deeper sites. Echinoderms were the most abundant phylum at each depth; ophiuroids accounted for 89% of individuals in photographs at 2000 m, echinoids were dominant at 3200 m (39%), and holothurians dominated at 4200 m (47%). We observed a 26% reduction in megafaunal density across the summit of the topographic rise relative to that documented on the continental slope at the same depth. However, the two communities at 3200 m were very similar in composition. Together, these data support the modified ‘archibenthal zone of transition’ framework for slope community patterns with distinct communities along the middle and lower slope (the upper slope was not evaluated here). This study fills a geographical gap by providing baseline information for a relatively pristine, high-latitude, deep-sea benthic ecosystem. As pressures grow for drilling, fishing and mining on high-latitude margins, such data can serve as a reference point for much-needed studies on the ecology, long-term dynamics, and anthropogenically induced change of these habitats.

Turbidity events observed in situ along the Congo submarine channel

Abstract As part of the multidisciplinary programme BIOZAIRE devoted to studying deep-sea benthic ecosystems in the Gulf of Guinea, particulate input and its relationship with near-bottom hydrodynamics were monitored using long-term moorings from 2000 to early 2005. Particular attention was given to material input through the Congo (ex-Zaire) submarine channel that extends 760 km from the Congo River mouth to the abyssal plain (>5100 m) near 6°S. Due to its direct connection to the Congo River, the Congo canyon and channel system are characterised by particularly active recent sediment transport. During this first in situ long-term monitoring along the channel, an energetic turbidity event was observed in January 2004 at three locations along the channel from 3420 to 4790 m in depth. This event tilted and displaced the moorings installed at 3420 m (site ZR′) and 4070 m (site ZD′), and resulted in high sediment deposition at all three mooring sites. The event moved at an average velocity of 3.5 m s−1 along the numerous channel meanders between 3420 and 4070 m, then at 0.7 m s−1 between 4070 m and the end of the channel at 4790 m. The particle cloud rose above the top of the valley at 4070 m (site ZD′), but not at 3420 m (site ZR′) where the channel was too deep. Lastly, the mooring line broke at site ZD′ in October 2004 probably due to a strong event like that of 2001 previously described by Khripounoff et al. [Khripounoff, A., Vangriesheim, A., Babonneau, N., Crassous, P., Denniellou, B., Savoye, B., 2003. Direct observation of intense turbidity activity in the Zaire submarine valley at 4000 m water depth. Marine Geology (194), 151–158]. Between these strong events, several peaks of high turbidity and particle flux occurred, but without noticeable current increases. These events were probably due to local sliding of sediment accumulated on the walls or terraces on the side of the channel. The area near 4000 m depth and the lobe appear to be the main depocentres of particulate input rich in organic matter derived from the Congo River.

Major viral impact on the functioning of benthic deep-sea ecosystems

Viruses are the most abundant biological organisms of the world's oceans. Viral infections are a substantial source of mortality in a range of organisms-including autotrophic and heterotrophic plankton-but their impact on the deep ocean and benthic biosphere is completely unknown. Here we report that viral production in deep-sea benthic ecosystems worldwide is extremely high, and that viral infections are responsible for the abatement of 80% of prokaryotic heterotrophic production. Virus-induced prokaryotic mortality increases with increasing water depth, and beneath a depth of 1,000 m nearly all of the prokaryotic heterotrophic production is transformed into organic detritus. The viral shunt, releasing on a global scale approximately 0.37-0.63 gigatonnes of carbon per year, is an essential source of labile organic detritus in the deep-sea ecosystems. This process sustains a high prokaryotic biomass and provides an important contribution to prokaryotic metabolism, allowing the system to cope with the severe organic resource limitation of deep-sea ecosystems. Our results indicate that viruses have an important role in global biogeochemical cycles, in deep-sea metabolism and the overall functioning of the largest ecosystem of our biosphere.

Response of an abyssal macrofaunal community to a phytodetrital pulse

MEPS Marine Ecology Progress Series Contact the journal Facebook Twitter RSS Mailing List Subscribe to our mailing list via Mailchimp HomeLatest VolumeAbout the JournalEditorsTheme Sections MEPS 355:73-84 (2008) - DOI: https://doi.org/10.3354/meps07240 Response of an abyssal macrofaunal community to a phytodetrital pulse A. K. Sweetman1,3,*, U. Witte2 1Max-Planck-Institut für Marine Mikrobiologie, Celsiusstrasse 1, 28359 Bremen, Germany 2Oceanlab, University of Aberdeen, Newburgh, Aberdeenshire AB41 6AA, UK 3Present address: Department of Oceanography, University of Hawaii at Manoa, 1000 Pope Road, Honolulu, Hawaii 96822, USA *Email: sweetman@hawaii.edu ABSTRACT: The majority of deep-sea benthic communities rely on particulate organic matter (POM) sinking from the euphotic zone for energy, much of which is delivered in pulsed events. But we know little about abyssal plain macrofaunal communities, their response to such events or their role in deep-sea carbon-cycling. In this study, we examined the composition of the macrofaunal community at Station M in the deep NE Pacific and assessed its short-term response to a simulated OM pulse in two 36 h in situ enrichment experiments. In each experiment, 1.2 gC m–2 of 13C-labelled Skeletonema costatum was deposited onto the seafloor using a benthic chamber lander. Macrofaunal abundance and biomass were significantly higher at 0 to 5 cm depth compared to 5 to 10 cm, and were dominated by the Nematoda and Crustacea, respectively. Twenty-five percent of the macrofauna specimens showed 13C signatures indicative of label ingestion, but specific uptake (Δδ13C) and C-turnover rates varied strongly between and within taxa. Two organisms, a single cumacean from 1 chamber and a paraonid polychaete from the second chamber, were responsible for the majority of C-uptake and had ingested up to 2.3% of their body weight in C. Macrofaunal C-turnover was much lower than recorded in the abyssal NE Atlantic, which is most likely due to differences in the timing of the experiments relative to the spring/summer bloom, different experimental durations and disparities in macrofaunal community structure. These results emphasize the degree of plasticity inherent in the macrofaunal response to a food pulse and stress the need for comprehensive in situ investigations to further our understanding of deep-sea benthic ecosystem functioning. KEY WORDS: Abyssal plain · Northeast Pacific · Deep sea · Pulse-chase experiments · δ13C · δ15N · Natural isotopic signatures · Benthic carbon remineralisation · Macrofauna Full text in pdf format PreviousNextCite this article as: Sweetman AK, Witte U (2008) Response of an abyssal macrofaunal community to a phytodetrital pulse. Mar Ecol Prog Ser 355:73-84. https://doi.org/10.3354/meps07240Export citation RSS - Facebook - Tweet - linkedIn Cited by Published in MEPS Vol. 355. Online publication date: February 26, 2008 Print ISSN: 0171-8630; Online ISSN: 1616-1599 Copyright © 2008 Inter-Research.

Long-term changes in microbial and biochemical parameters in the Central Indian Basin

Natural kilometer-scale spatial variability and seasonal variability in deep-sea sediments at ∼5000 m depth was studied at two reference sites in the Central Indian Basin. Biochemical parameters such as the labile organic matter (LOM) (consisting of carbohydrates, protein and lipids) and total organic matter (TOM) and biological parameters such as total counts of bacteria and adenosine triphosphate (ATP) were estimated after an interval of 54 months. These were compared with nine locations (12–17 km away) where an artificial sediment disturbance was created by a hydraulic benthic disturber. Besides the above-mentioned parameters, extracellular enzymes, alkaline phosphatase, aminopeptidase and lipase were also monitored at these nine locations. Post-disturbance (PoD1) observations were carried out immediately after the benthic disturbance experiment and compared with the pre-disturbance (PreD) pattern. The restoration process in the test site was studied after a period of 44 months (PoD2). There was a drastic reduction in total counts of bacteria, ATP, carbohydrates and lipids accompanied by a two- to four-fold increase in protein content at the undisturbed reference sites after 54 months. A similar reduction in these parameters was observed in the PoD1 compared to the PreD stage at the test site. An initial increase in TOM content and in the activities of three extracellular enzymes in the PoD1 stage was followed by a reduction in TOM, ATP and organic P content and activities of the extracellular enzymes in the PoD2 phase (i.e. after 44 months). Reduction in ATP levels suggests a negative impact of resedimentation on living biomass in the sediments. Although LOM, protein concentrations and the protein/carbohydrate ratio increased in the PoD2 phase relative to the PoD1 phase, the bacterial numbers were below the concentrations in the PreD stage, indicating slow restoration of benthic conditions. Long-term analyses of the indicator parameters thus showed mixed effects of sediment disturbance. A similar pattern at the undisturbed reference location suggests that natural (seasonal) variability outweighed artificial variability caused by the benthic disturbance in the Central Indian Basin and the former is responsible for bringing about changes in deep-sea benthic ecosystem on long-time scales.

Mass deposition of jellyfish in the deep Arabian Sea

In December 2002 large numbers of dead jellyfish, Crambionella orsini (Vanhoffen 1888), were observed on the seabed over a wide area of the Arabian Sea off the coast of Oman, at depths between 350 and 3300 m. Moribund jellyfish were seen tumbling down the continental slope. Large aggregations of dead jellyfish were evident within canyons and on the continental rise. At the deepest stations, patches of rotting, coagulated jellyfish occurred. The patches were several metres in diameter, at least 7 cm thick and covered about 17% of the sediment surface. At other locations on the continental rise the seafloor was covered in a thin, almost continuous, layer of jelly “slime,” a few millimetres thick, or was littered with individual jellyfish corpses. Photographic transects were used to estimate the amount of carbon associated with the jelly detritus. The standing stock of carbon varied between 1.5 and 78 g C m-2, the higher figure exceeding the annual downward flux of organic carbon, as measured by sediment traps, by more than an order of magnitude. The episodic nature of jellyfish blooms, which may be modulated by global change phenomena, provides a hitherto unknown mechanism for large-scale spatial and temporal patchiness in deep-sea benthic ecosystems.

In situ Enclosure Experiment Using a Benthic Chamber System to Assess the Effect of High Concentration of CO2 on Deep-Sea Benthic Communities

In order to evaluate the environmental impact associated with sequestration of carbon dioxide in the deep sea, a free fall type field experimental device, the benthic chamber, was developed. In situ experiments to expose deep-sea communities to elevated concentrations of carbon dioxide (average of 20,000 ppm, 5,000 ppm and control) were carried out using this device 3 times, viz., in the winter of 2002 and in the spring and the summer of 2003, in the Kumano Trough at a depth of 2,000 m. In the long-term experiments (about two weeks in winter of 2002 and summer of 2003), the abundance of meiobenthos declined whereas that of bacteria increased under the condition of 20,000 ppm carbon dioxide compared with the control. Among meiofauna, the abundance of foraminifers at the same concentration of carbon dioxide became less than the control even in the short-term (3 days in spring of 2003) experiment, suggesting that organisms with a calcium carbonate exoskeleton are more sensitive to the raised concentration of carbon dioxide. The respiration rate of the benthic community exposed to 20,000 ppm was lower in the early stage of the experiment than in the latter half, whereas it was opposite under the condition of 5,000 ppm. The increase of biological activity in the 20,000 ppm exposure group is probably due to an increase of bacteria adapted to high carbon dioxide concentrations. The present results suggest that the influence of carbon dioxide on the deep-sea benthic ecosystem does not follow a simple, linear relationship with concentration.

Mapping deep‐water gas emissions with sidescan sonar

Emissions of methane gas from cold seeps on the seafloor have a strong impact on a number of biogeochemical processes. These processes include the development of deepsea benthic ecosystems via the process of anaerobic oxidation of methane [Boetius et al., 2000] or the precipitation of carbonates [Ritger et al., 1987]. The fluxes of other chemical species associated with methane emissions may even influence the chemical composition of seawater [Aloisi et al., 2004]. Such gas emissions may have been much more intensive in the past with a strong impact on global climate [Dickens, 1999], as suggested by carbon isotope data.

GIANT BATHYSIPHON (ASTRORHIZINA: FORAMINIFERA) FROM THE LATE CRETACEOUS HUNTERS COVE FORMATION, SOUTHWESTERN OREGON

ORAMINIFERANS BELONGING to the genus Bathysiphon G. O. and M. Sars in G. O. Sars, 1872 are among the largest fossil protists ever described, some species having tubular tests exceeding 100 mm in length (Miller, 1995). One species, Bathysiphon aaltoi Miller, 1986, from mid-Cretaceous fine-grained turbidites within the Franciscan Complex of northern California, may be the largest fossil foraminiferan ever collected with an estimated test length of approximately 150 mm (Miller, 1991, 1997). Species of comparable size are known from the modern deep-ocean floor (where they often rival or surpass the size of co-occurring metazoans) and are an important component of deep-sea benthic ecosystems (Gooday, 1983; Gage and Tyler, 1991; Gooday et al., 1992). In the tectonically active parts of the Pacific Rim, these giant protists are commonly found in flysch successions in association with typical deepwater trace fossils such as graphoglyptids. These remarkable tubular fossils often are the only body fossils found in abundance in Mesozoic to Cenozoic turbidite deposits in these regions, and the association appears to be indicative of deep seafloor settings strongly influenced by sedimentation and nutrient cycling from adjacent landmasses. In the regional geologic literature, however, bathysiphonids have been misidentified as body fossils of ‘‘worms’’ or as tubular trace fossils. In this report, I describe a new species of Bathysiphon from the Hunters Cove Formation, a thick sequence of Campanian‐ Maastrichtian turbidites in southwestern Oregon (Figs. 1, 2). This new taxon joins the pantheon of largest foraminiferans on record, having glassy, tubular tests up to 100 mm long and 5 mm wide. It differs from other large, Cretaceous Bathysiphon in having straight, thin-walled tests featuring vague axial striae on the outer surface. The taphonomy and paleoecology of this giant foraminiferan are briefly evaluated based on comparison with modern examples of Bathysiphon and with other Cretaceous examples from the same region. GEOLOGIC SETTING

The deep-sea floor ecosystem: current status and prospects of anthropogenic change by the year 2025

The goal of this paper is to review current impacts of human activities on the deep-sea floor ecosystem, and to predict anthropogenic changes to this ecosystem by the year 2025. The deep-sea floor ecosystem is one of the largest on the planet, covering roughly 60% of the Earth's solid surface. Despite this vast size, our knowledge of the deep sea is poor relative to other marine ecosystems, and future human threats are difficult to predict. Low productivity, low physical energy, low biological rates, and the vastness of the soft-sediment deep sea create an unusual suite of conservation challenges relative to shallow water. The numerous, but widely spaced, island habitats of the deep ocean (for example seamounts, hydrothermal vents and submarine canyons) differ from typical deep-sea soft sediments in substrate type (hard) and levels of productivity (often high); these habitats will respond differently to anthropogenic impacts and climate change. The principal human threats to the deep sea are the disposal of wastes (structures, radioactive wastes, munitions and carbon dioxide), deep-sea fishing, oil and gas extraction, marine mineral extraction, and climate change. Current international regulations prohibit deep-sea dumping of structures, radioactive waste and munitions. Future disposal activities that could be significant by 2025 include deep-sea carbon-dioxide sequestration, sewage-sludge emplacement and dredge-spoil disposal. As fish stocks dwindle in the upper ocean, deep-sea fisheries are increasingly targeted. Most (perhaps all) of these deep-sea fisheries are not sustainable in the long term given current management practices; deep-sea fish are long-lived, slow growing and very slow to recruit in the face of sustained fishing pressure. Oil and gas exploitation has begun, and will continue, in deep water, creating significant localized impacts resulting mainly from accumulation of contaminated drill cuttings. Marine mineral extraction, in particular manganese nodule mining, represents one of the most significant conservation challenges in the deep sea. The vast spatial scales planned for nodule mining dwarf other potential direct human impacts. Nodule-mining disturbance will likely affect tens to hundreds of thousands of square kilometres with ecosystem recovery requiring many decades to millions of years (for nodule regrowth). Limited knowledge of the taxonomy, species structure, biogeography and basic natural history of deep-sea animals prevents accurate assessment of the risk of species extinctions from large-scale mining. While there are close linkages between benthic, pelagic and climatic processes, it is difficult to predict the impact of climate change on deep-sea benthic ecosystems; it is certain, however, that changes in primary production in surface waters will alter the standing stocks in the food-limited, deep-sea benthic. Long time-series studies from the abyssal North Pacific and North Atlantic suggest that even seemingly stable deep-sea ecosystems may exhibit change in key ecological parameters on decadal time scales. The causes of these decadal changes remain enigmatic. Compared to the rest of the planet, the bulk of the deep sea will probably remain relatively unimpacted by human activities and climate change in the year 2025. However, increased pressure on terrestrial resources will certainly lead to an expansion of direct human activities in the deep sea, and to direct and indirect environmental impacts. Because so little is known about this remote environment, the deep-sea ecosystem may well be substantially modified before its natural state is fully understood.

Biological responses to seasonally varying fluxes of organic matter to the ocean floor: a review

Deep-sea benthic ecosystems are sustained largely by organic matter settling from the euphotic zone. These fluxes usually have a more or less well-defined seasonal component, often with two peaks, one in spring/early summer, the other later in the year. Long time-series datasets suggest that inter-annual variability in the intensity, timing and composition of flux maxima is normal. The settling material may form a deposit of phytodetritus on the deep-seafloor. These deposits, which are most common in temperate and high latitude regions, particularly the North Atlantic, evoke a response by the benthic biota. Much of our knowledge of these responses comes from a few time-series programmes, which suggest that the nature of the response varies in different oceanographic settings. In particular, there are contrasts between seasonal processes in oligotrophic, central oceanic areas and those along eutrophic continental margins. In the former, it is mainly small organisms (bacteria and protozoans) that respond to pulsed inputs. Initial responses are biochemical (e.g. secretion of bacterial exoenzymes) and any biomass increases are time lagged. Increased metabolic activity of small organisms probably leads to seasonal fluctuations in sediment community oxygen consumption, reported mainly in the North Pacific. Metazoan meiofauna are generally less responsive than protozoans (foraminifera), although seasonal increases in abundance and body size have been reported. Measurable population responses by macrofauna and megafauna are less common and confined largely to continental margins. In addition, seasonally synchronised reproduction and larval settlement occur in some larger animals, again mainly in continental margin settings. Although seasonal benthic responses to pulsed food inputs are apparently widespread on the ocean floor, they are not ubiquitous. Most deep-sea species are not seasonal breeders and there are probably large areas, particularly at abyssal depths, where biological process rates are fairly uniform over time. As with other aspects of deep-sea ecology, temporal processes cannot be encapsulated by a single paradigm. Further long time-series studies are needed to understand better the nature and extent of seasonality in deep-sea benthic ecosystems.

Deep-sea ostracode species diversity: response to late Quaternary climate change

Late Quaternary ostracode assemblages from the North Atlantic Ocean were studied to establish the effect of climatic changes of the past 210,000 yr (marine oxygen isotope stages 7–1) on deep-sea benthic biodiversity and faunal composition. Two-hundred and twenty five samples from the Chain 82-24 Core 4PC (41°43′N, 32°51′W, 3427 m water depth) on the western Mid-Atlantic Ridge revealed high amplitude fluctuations in ostracode abundance and diversity coincident with orbital and suborbital scale climate oscillations measured by several paleoceanographic proxy records. During the past 210,000 yr, ostracode biodiversity as measured by species number ( S) and the Shannon–Weaver index, H( S), oscillated from H( S)=0.4 during glacial periods (marine isotope stages 6, 5d, 5b, 4, and 2) to H( S)=1.1 during interglacial and interstadial periods (stages 7, 5e, 5c, 5a, 3 and 1). A total of 23 diversity peaks could be recognized. Eleven of these signify major periods of high diversity [ H( S)>0.8, S = 10–21] occurring every 15–20 ka. Twelve were minor peaks which may represent millennial-scale diversity oscillations. The composition of ostracode assemblages varies with Krithe-dominated assemblages characterizing glacial intervals, and Argilloecia–Cytheropteron characterizing deglacials, and trachyleberid genera ( Poseidonamicus, Echinocythereis, Henryhowella, Oxycythereis) abundant during interglacials. Diversity and faunal composition changes can be matched to independent deep-sea paleoceanographic tracers such as benthic foraminiferal carbon isotopes, Krithe trace elements (Mg/Ca ratios), and to North Atlantic region climate records such as Greenland ice cores. When interpreted in light of ostracode species' ecology, these faunal and diversity patterns provide evidence that deep-sea benthic ecosystems experience significant reorganization in response to climate changes over orbital to millennial timescales.

210Po as a Tracer of Variations of Trophic Input to Deep-sea Benthic Ecosystems: A Study of the Deep-sea Amphipod Eurythenes Gryllus from the Tropical Atlantic

Deep-sea amphipods were trapped at different heights above the sea floor (4500 m depth) during two periods (February and September 1991) in the tropical Atlantic Ocean off Mauritania, and were analysed for their 210 Po content. In fact, 210 Po variations were considered an adequate tracer for reflecting changes in trophic inputs at this location. 210 Po and 210 Pb concentrations ranged from 23 to 286 Bq.kg -1 dry weight and from 12 to 67 Bq.kg -1 dry weight, respectively, when considering the whole organisms with no clear trend with the height of capture above the sea floor. A comparison with 210 Po concentrations in Eurythenes gryllus and other crustaceans from different locations in the Atlantic was carried out. 210 Po concentrations in deep-sea amphipods appeared far less important and could be related to their peculiar feeding strategy. A significant inverse correlation was found between 210 Po concentrations and weight of the amphipods for the two seasons. Allometric equations for 210 Po concentrations revealed similar slopes for the two periods of sampling; however, 210 Po concentrations were higher in September compared to February. This difference could be linked to temporal variations both in the mass and organic carbon fluxes into this deep-sea environment.

The Biology of Deep-Sea Foraminifera: A Review of Some Advances and Their Applications in Paleoceanography

Foraminifera commonly dominate ocean-floor eukaryotic communities. They also are the most abundant benthic organisms to be preserved in the post-Paleozoic deep-sea fossil record. Living faunas contain numerous delicate, soft-bodied agglutinated forms which have virtually no fossilization potential, whereas fossil faunas usually consist of calcareous and more robust agglutinated taxa. Recent observations have emphasized the important, often active role that foraminifera play in the dynamics and structuring of deep-sea benthic ecosystems. Foraminiferal abundances are closely linked to levels of organic matter input, and to dissolved oxygen concentrations in the near-bottom water which, in part, are related inversely to the magnitude of organic fluxes

Responses by benthic organisms to inputs of organic material to the ocean floor: a review

Most of the photosynthetically produced organic material reaching the ocean-floor is transported as settling particles, among which larger particles such as faecal pellets and macroaggregates (marine snow) are particularly important. Recent studies in the northeastern Atlantic have demonstrated that macroaggregates originating from the euphotic zone settle at a rate of approximately 100-150 m d -1 to form a deposit (phytodetritus) on the sediment surface. Bacteria and protozoa (flagellates and foraminifers) rapidly colonize and multiply on phytodetritus, while large deposit feeding animals ingest it. Other inputs, for example Sargassum , wood and vertebrate carcasses, also evoke a rapid response by benthic organisms. However, the taxa that respond depend on the form of the organic material. The intermittent or seasonally pulsed nature of phytodetritus and many other inputs regulate the population dynamics and reproductive cycles of some responding species. These are often opportunists that are able to utilize ephemeral food resources and, therefore, undergo rapid fluctuations in population density. In addition, the patchy distribution of much of the organic material deposited on the ocean-floor probably plays a major role in structuring deep-sea benthic ecosystems.

Trophic strategies in carnivorous fishes: their significance in energy transfer in the deep-sea benthic ecosystem (Meriadzek Terrace — Bay of Biscay)

Trophic relationships between fish species have been analyzed as a contribution to the construction of an energy transfer model between various groups in the benthic ecosystem of Meriadzek Terrace, located at 170 miles off the west French coast at a depth of 2100m. Attention was given to the carnivorous megafaunal compartment. The importance of the carnivore megafaunal biomass and its part in energy transfer is still unresolved. On the Meriadzek Terrace, numerous Chondrichthyes (sharks and chimeroids) were frequently observed from the manned submersible Cyana or by baited camera; their biomass appears to be exceptionally large. Results of a photographic survey conducted using the unmanned free vehicle Epaulard have given estimates of 22 per 10 4m 2 of teleost fishes (belonging to 8 different families and one suborder) but included no Chondrichthyes (except one specimen of Rajidae). From baited camera observations, however, we conclude that carcasses falling to the bottom on the Terrace are exploited only by selacians as the bait was never seen to be eaten by macrourids, morids or synaphobranchids although they always attended the baits before the arrival of large scavengers. Trophic specialization strategies occur in this deep-sea ecosystem and our further studies on the energy transfer model for the Meriadzek Terrace will differentiate between strategies exploiting two sources of organic input to the bottom environment: organic particles and large carcasses. In one case, carbon transfer is via the whole benthic food web in which each link is specified by size and trophic behaviour. This pathway ends in the carnivorous megafaunal compartment, which includes benthic fishes, consumers of the local epibenthic and sometimes inbenthic, macro- and megafauna. The second pathway originates with carcass falls, and carbon transfer is direct to the benthopelagic scavengers observed frequently at the Meriadzek Terrace.