John D. Gage

Monitoring environmental variability around cold-water coral reefs: the use of a benthic photolander and the potential of seafloor observatories

The environmental sensitivies of cold-water corals and their associated biota are likely to be determined by the natural variability of the cold-water coral reef environment. The sensitivity of reef biota to sedimentation and resuspension events is largely unknown and the influence of seasonal phytodetrital deposition is poorly understood. Here we describe the use of a benthic photolander to monitor this variability by the Sula Ridge reef complex on the mid-Norwegian continental shelf and from the Galway carbonate mound in the Porcupine Seabight. The photolander provides a platform for time-lapse digital and film cameras to image the seabed while recording the current regime and optical characteristics (light transmission, backscatter and fluorescence) of the seawater. In its first two deployments carried out in 2001 and 2002 by the Sula Ridge the lander recorded a dynamic environment around the reef site with a tidal current regime and periods of sediment resuspension. Current speeds by the Sula Ridge reef complex reached a maximum of 28 cm s−1 and 70 cm s−1 on the Galway carbonate mound, reinforcing much speculation about the dependence of these communities on current-swept conditions. Seabed photographs show intense feeding activity of echiuran worms (Bonellia viridis) near the Sula Ridge reef complex pointing to rapid bioturbation of the sediment. Fish were recorded sheltering near sponges that had colonised glacial dropstones. Longer term monitoring in situ is needed for study of seasonal change, to identify functional roles of associated fauna and to monitor potential coral spawning events. Benthic landers and seafloor observatories have great potential in these areas. Only with a better understanding of the natural variability of the cold-water coral environment can informed decisions about the environmental sensitivity of cold-water coral reefs and their management be made.

Why are there so many species in deep-sea sediments?

Abstract High species diversity in samples of macrobenthos of deep-sea sediments is now well established. But a consensus on the processes regulating this unexpectedly species-rich coexistence in metazoan species at the deep-sea bed is still elusive. This review takes a broad approach by examining differences between marine and terrestrial biodiversity in the context of the following: scale of sampling effort on which our knowledge is based; the species concept as applied in the past to taxonomic studies on deep-sea benthic organisms; scaling differences and size related patterns in community structure and habitat complexity and differences in potential for co-evolution. Latitudinal and bathymetric patterns are summarised in relation to habitat variability and distributional range and in relation to J.S. Grays (1994) claim that benthic diversity may be equally high on the continental shelf. Evidence is first reviewed for neo-Darwinian competitive co-existence based on niche specialisation and habitat partitioning. Although evidence for any dietary specialisation is sparse, biologically generated heterogeneity in the sediment, such as from mounds, burrows, tubes and ‘mud balls’, will persist longer and contribute more to niche diversification than in shallow water owing to slower obliteration by sedimentation and water turbulence. Second, the role of periodic small-scale disturbance in creating a shifting dynamic of invasion and species succession in patches of new, open habitat, is reviewed in relation to data from sea bed observations and experiments. Such disturbances may arise from effects ranging from organic enrichment as falls of carcases or dead plants, seaweed or wood, to patchy phytodetrital deposition from the surface. Environmental patchiness of both kinds may be important in determining small scale, predominantly non-competitive co-existence, particularly in a habitat lacking large-scale isolating barriers and open to chance, low-intensity recruitment from propagules of a wide taxonomic and functional spectrum of biota. Disturbance may also be expressed as diffuse, large-scale effects caused by sea bed currents. At their most extreme, ‘benthic storms’ erode and re-deposit sediment and create a regime of intermittent disturbance that reduces species richness; but more moderate flow may enhance recruitment and remould the small-scale sediment landscape to create new habitat. The wide range in habitat partitioning and disturbance processes that may be involved in controlling species richness make realistic modelling studies challenging. Careful description of the full spatio-temporal range in environmental heterogeneity at the deep-sea bed is badly needed. Further manipulative experiments, and perhaps pilot ‘industrial’-scale studies involving laying bulky inert or organically enriched waste on the bed, will certainly also be useful in improving our knowledge of processes controlling deep-sea macrobenthic species richness.

The cold-water coral Lophelia pertusa (Scleractinia) and enigmatic seabed mounds along the north-east Atlantic margin: are they related?

In this study, an updated distribution of Lophelia pertusa between the Porcupine Seabight and Norwegian shelf is presented. It seems unlikely that enigmatic mound structures observed at water depths of more than 570 m during acoustic seabed surveys, particularly to the west of the Shetland Islands, are related to the occurrence of L. pertusa. At these depths in the Faroe-Shetland Channel, the predominant influence of cold Arctic water precludes its growth. Iceberg dumpsites are also considered unlikely explanations for the origin of these mounds, and they are interpreted as most likely to be related to the release of fluids at the seabed. When mound structures were investigated, no scleractinian corals were recovered at water depths >500 m. This study shows the importance of seabed temperature as an environmental control on cold-water coral distribution. The significance of cold-water coral habitats in sustaining high levels of local-scale biodiversity is now becoming apparent in parallel with increased hydrocarbon extraction and fishing activity beyond the shelf edge. There is growing evidence that these areas have been marked by the passage of deep-water trawls. It seems likely that trawling activity has already reduced the extent of cold-water coral distribution in this region of the north-east Atlantic.

Patterns in polychaete abundance and diversity from the Madeira Abyssal Plain, northeast Atlantic

Polychaete abundance and diversity patterns from the Madeira Abyssal Plain (MAP) were studied together with data from three other sites on the northeastern Atlantic abyssal plains. Polychaete abundance at MAP was significantly lower than at any of the other sites, including those lying under comparable productivity regimes. Analysis of diversity, using rarefaction and species counts per unit area, suggests that MAP is extremely species poor and shows dominance by a few common species. The MAP site is characterised by a superficial layer of turbidite sediment, and the hypothesis is put forward that the unusual sediment characteristics at MAP have affected macrofaunal abundance over a vast area (>2000 km2). Analysis of species composition indicates that the MAP site is not faunistically unique; rather it contains a high proportion of widespread, abundant, cosmopolitan species. We suggest that these are the opportunists of the abyssal benthic habitat. Differences in abundance between the other abyssal sites are the result of both productivity and local environmental conditions. Equitability at the other north Atlantic sites is not affected by productivity, although the actual number of species per unit area is affected, showing a south–north gradient.

A Preliminary Survey of the Benthic Macrofauna and Sediments in Lochs Etive and Creran, Sea-Lochs Along the West Coast of Scotland

The hydrographic conditions in the two sea-lochs is briefly described where relevant to the survey of the benthos. Seasonal changes in temperature and salinity in Loch Etive appear to follow roughly the local coastal pattern but usually with a lag in timing that is considered to be because of the restricting effect of the twin sills. Replenishment of deep-basin water by ventilation over the sills is spasmodic and controlled by surface stratification; the latter being due primarily to freshwater runoff. This situation contrasts with Loch Creran which has only a small catchment and where the seasonal hydrography closely follows the pattern outside. Other possible reasons for the differences observed between the two lochs are discussed. The bottom fauna of both lochs was sampled semi-quantitatively with an anchor dredge at 22 stations in Loch Etive and 8 in Loch Creran at depths from 7–117 m. A detailed list of the fully marine fauna present is given. The species occurrences in the single-haul samples are compared objectively using a coefficient of similarity of the presence/absence records. An attempt is made to relate the groupings shown up to the classical ‘bottom community’ concept, to the feeding-type categories of the bottom animals and to analysis of particle size of sediment samples taken at the same stations. A correlation to sediment type was apparent. This is considered to owe as much to local hydrographic conditions which the sediment reflect as to the sediment itself.

Seabed photography, environmental assessment and evidence for deep-water trawling on the continental margin west of the Hebrides

A photographic survey in 1998 of the seabed along depth transects from 700 to 1300 m across the N.E. Atlantic continental slope off north-west Scotland shows clear depth-related change in sediment type and megabenthic community in an environment where biological communities and species distributions are poorly known. Small-scale features, such as trawl marks and dense fields of xenophyophores, were resolved that may have remained unknown using conventional sampling or lower resolution imaging techniques. Because xenophyophores accumulate barite, a constituent of some drilling muds, their local-scale occurrences will be important to baseline environmental survey prior to hydrocarbon prospecting in deep water. Our results indicate that deep-sea trawling is physically impacting the seabed to depths of more than 1000 m. The persistence and biological consequence of this impact is unknown, but may depend on sediment type and natural physical disturbance. Comparison with similar seabed photographs taken from a neighbouring area in 1988, which show a high incidence of trawl marks, indicates that such impacts have been taking place over at least 10 years.

Deep-sea benthic community and environmental impact assessment at the Atlantic Frontier

Abstract The seabed community provides a sensitive litmus for environmental change. North Sea analysis of benthic populations provides an effective means for monitoring impacts from mans interventions, such as offshore oil exploitation and fishing, against baseline knowledge of the environment. Comparable knowledge of the benthic biology in the deep waters of the Atlantic Frontier beyond the N.E. Atlantic shelf edge is poorly developed. But uncertainties should not encourage assumptions and extrapolations from the better-known conditions on the continental shelf. While sampling at present still provides the best means to assess the health of the deepwater benthic habitat, protocols developed for deep-sea fauna should be applied. These are necessary because of (a) lower faunal densities, (b) higher species richness, (c) smaller body size, and (d) to ensure comparability with other deep-sea data. As in the North Sea, species richness and relative abundance can be analysed from quantitative samples in order to detect impacts. But analysis based on taxonomic sufficiency above species level is premature, even if arguably possible for coastal communities. Measures also need to ensure identifications are not forced to more familiar coastal species without proper study. Species-level analysis may be applied to seabed photographs of megafauna in relation to data on bottom environment, such as currents and the sediment, to monitor the health of the deep-water community. Although the composition of higher taxa in the benthic community is broadly similar to soft sediments on the shelf, concordance in sensitivities is speculative. Moreover, new organisms occur, such as giant protozoan xenophyophores, unknown on the continental shelf, whose sensitivities remain conjectural. Past knowledge of the benthic biology of the deep-water areas off Scotland is based on scattered stations and some more focussed, multidisciplinary studies, and should be significantly augmented by the results from the oil industry-funded Atlantic Margin Environmental Study cruises in 1996 and 1998. A predominantly depth-related pattern in variability applies here as found elsewhere in the deep ocean, and just sufficient knowledge-based predictive power exists to make comprehensive, high-resolution grid surveys unnecessary for the purpose of broad-scale environmental assessment. But new, small-scale site surveys remain necessary because of local-scale variability. Site survey should be undertaken in the context of existing knowledge of the deep sea in the UK area of the Atlantic Frontier and beyond, and can itself usefully be structured as tests of a projection from the regional scale to reduce sampling effort. It is to the benefit of all stakeholders that environmental assessment aspires to the highest scientific standards and contributes meaningfully to context knowledge. By doing so it will reduce uncertainties in future impact assessments and hence contribute usefully to environmental risk management.

Patterns in deep-sea macrobenthos at the continental margin: standing crop, diversity and faunal change on the continental slope off Scotland

Depth-related patterns of macrobenthic community structure and composition have been studied from box-core samples from the Scottish continental slope where deep-sea trawling and oil exploration are becoming increasingly important. There is a strong pattern of declining biomass and faunal abundance with increasing depth, but results also indicate reduced biomass and numbers of macrobenthos in the shallowest samples from just below the shelf edge where there are coarse sediments and a regime of strong bottom currents. There is also reduced species diversity at the shallowest stations, probably caused by hydrodynamic disturbance, but no clear mid-slope peak in species diversity as described from the northwest Atlantic. Taxonomic composition of the macrobenthic community shows most change between about 1000 and 1200 m, expressed as a major dichotomy in multivariate analysis by cluster analysis and ordination. It also shows up as a step-like increase in the rate of accumulation of new macrofaunal species. This corresponds to a change in hydrodynamic regime, from a seabed rich in suspension- and interface-feeding epifauna, to one where biogenic traces from large, burrowing deposit feeders are well developed, and visible epifauna rare in seabed photographs. It also corresponds to the depth zone where earlier study of megafaunal echinoderms in trawl and epibenthic sled samples also shows a clear peak in across-slope rate of change in faunal composition.

Evidence for coupling the vertical flux of phytodetritus to the diet and seasonal life history of the deep-sea echinoid Echinus affinis

Abstract Data from a temporal sampling programme at 2200 m depth provide evidence for a relationship between the diet of the deep-sea echinoid Echinus affinis and deposition of phytodetritus. Nonlinear regression and “lack of fit” analysis resulted in the identification of a number of seasonal fluctuations in the organ and gut content indices of E. affinis . The amount of gut contents were low in winter and high during spring and summer months. The proportionof organic matter in the gut contents, however, did not show a significant seasonal variation. Planktonic and benthic material was identified from the gut contents using electron microscopy. All these data support the hypothesis that the seasonal deposition of phytodetritus provides energy for growth and reproduction.

Vertical distribution correlates with pressure tolerances of early embryos in the deep-sea asteroid Plutonaster bifrons

The astropectinid asteroid Plutonaster bifrons (Wyville Thomson) occurs on the continental slope of the north-east Atlantic between 1000 and 2500 m depths. As in most deepsea animals, the factors limiting bathymetric distribution of this species are unknown. Eggs were fertilized in vitro and incubated through the early embryonic cleavage stages at pressures that correspond to depths from 0 to 3000 m. The highest percentage of normal development occurred near the peak of the species distribution (2000 m), and virtually no normal development occurred at a pressure corresponding to 3000 m depth. Develop-mental rate was retarded at pressures higher and lower than those found near 2000 m. These experiments indicate that embryonic pressure tolerances could determine both the upper and lower bathymetric limits of distribution for this species.