Brian J. Bett

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.

Long-term change in the megabenthos of the Porcupine Abyssal Plain (NE Atlantic)

A radical change in the abundance of invertebrate megafauna on the Porcupine Abyssal Plain is reported over a period of 10 years (1989–1999). Actiniarians, annelids, pycnogonids, tunicates, ophiuroids and holothurians increased significantly in abundance. However, there was no significant change in wet weight biomass. Two holothurian species, Amperima rosea and Ellipinion molle, increased in abundance by more than two orders of magnitude. Samples from the Porcupine Abyssal Plain over a longer period (1977–1999) show that prior to 1996 these holothurian species were always a minor component of the megafauna. From 1996 to 1999 A. rosea was abundant over a wide area of the Porcupine Abyssal Plain indicating that the phenomenon was not a localised event. Several dominant holothurian species show a distinct trend in decreasing body size over the study period. The changes in megafauna abundance may be related to environmental forcing (food supply) rather than to localised stochastic population variations. Inter-annual variability and long-term trends in organic matter supply to the seabed may be responsible for the observed changes in abundance, species dominance and size distributions.

Material supply to the abyssal seafloor in the northeast Atlantic

Downward particle flux was measured using sediment traps at various depths over the Porcupine Abyssal Plain (water depth ~4850 m) for prolonged periods from 1989 to 1999. A strong seasonal pattern of flux was evident reaching a maximum in mid-summer. The composition of the material changed with depth, reflecting the processes of remineralisation and dissolution as the material sank through the water column. However, there was surprisingly little seasonal variation in its composition to reflect changes in the biology of the euphotic zone. Currents at the site have a strong tidal component with speeds almost always less than 15 cm/sec. In the deeper part of the water column they tend to be northerly in direction, when averaged over periods of several months. A model of upper ocean biogeochemistry forced by meteorology was run for the decade in order to provide an estimate of flux at 3000 m depth. Agreement with measured organic carbon flux is good, both in terms of the timings of the annual peaks and in the integrated annual flux. Interannual variations in the integrated flux are of similar magnitude for both the model output and sediment trap measurements, but there is no significant relationship between these two sets of estimates. No long-term trend in flux is evident, either from the model, or from the measurements. During two spring/summer periods, the marine snow concentration in the water column was assessed by time-lapse photography and showed a strong peak at the start of the downward pulse of material at 3000 m. This emphasises the importance of large particles during periods of maximum flux and at the start of flux peaks. Time lapse photographs of the seabed show a seasonal cycle of coverage of phytodetrital material, in agreement with the model output both in terms of timing and magnitude of coverage prior to 1996. However, after a change in the structure of the benthic community in 1996 no phytodetritus was evident on the seabed. The model output shows only a single peak in flux each year, whereas the measured data usually indicated a double peak. It is concluded that the observed double peak may be a reflection of lowered sediment trap efficiency when flux is very high and is dominated by large marine snow particles. Resuspension into the trap 100 m above the seabed, when compared to the primary flux at 3000 m depth (1800 mab) was lower during periods of high primary flux probably because of a reduction in the height of resuspension when the material is fresh. At 2 mab, the picture is more complex with resuspension being enhanced during the periods of higher flux in 1997, which is consistent with this hypothesis. However there was rather little relationship to flux at 3000 m in 1998. At 3000 m depth, the Flux Stability Index (FSI), which provides a measure of the constancy of the seasonal cycle of flux, exhibited an inverse relationship with flux, such that the highest flux of organic carbon was recorded during the year with the greatest seasonal variation.

Meiobenthos of the deep Northeast Atlantic

This chapter throws the attention on the meiobenthos of the deep northeast Atlantic. The main purpose of this chapter is to summarize new results from an area lying between 15°N and 53°N and extending from the continental margin of western Europe and northwest Africa to the Mid-Atlantic Ridge. It considers first the nature and scope of meiofaunal research in the northeast Atlantic and then discuss the environmental parameters, which are believed to influence meiofaunal organisms. This chapter then discusses the various types and scales of pattern observed among meiofaunal populations within the study area, progressing from the large-scale bathymetric and latitudinal trends and then to small-scale horizontal patterns within particular areas. Faunal densities and faunal composition are considered separately and compared with data from other regions. This chapter also deals with the distribution of meiofauna within sediment profiles and the temporal variability of populations. This chapter concludes by discussing the recent review of deep-sea meiofauna, which focused mainly on the abundance and biomass data from different oceans and on the relationship between the biomass of the meiofauna and that of other faunal components

Climate, carbon cycling, and deep-ocean ecosystems

Climate variation affects surface ocean processes and the production of organic carbon, which ultimately comprises the primary food supply to the deep-sea ecosystems that occupy ≈60% of the Earths surface. Warming trends in atmospheric and upper ocean temperatures, attributed to anthropogenic influence, have occurred over the past four decades. Changes in upper ocean temperature influence stratification and can affect the availability of nutrients for phytoplankton production. Global warming has been predicted to intensify stratification and reduce vertical mixing. Research also suggests that such reduced mixing will enhance variability in primary production and carbon export flux to the deep sea. The dependence of deep-sea communities on surface water production has raised important questions about how climate change will affect carbon cycling and deep-ocean ecosystem function. Recently, unprecedented time-series studies conducted over the past two decades in the North Pacific and the North Atlantic at >4,000-m depth have revealed unexpectedly large changes in deep-ocean ecosystems significantly correlated to climate-driven changes in the surface ocean that can impact the global carbon cycle. Climate-driven variation affects oceanic communities from surface waters to the much-overlooked deep sea and will have impacts on the global carbon cycle. Data from these two widely separated areas of the deep ocean provide compelling evidence that changes in climate can readily influence deep-sea processes. However, the limited geographic coverage of these existing time-series studies stresses the importance of developing a more global effort to monitor deep-sea ecosystems under modern conditions of rapidly changing climate.

Temporal variability in phytodetritus and megabenthic activity at the seabed in the deep Northeast Atlantic

We report a ten-year study of the abundance and activity of megabenthos on the Porcupine Abyssal Plain, northeast Atlantic, together with observations on the occurrence of phytodetritus at the deep-sea floor (4850 m). Using the Southampton Oceanography Centre time-lapse camera system, ‘Bathysnap’, we have recorded a radical change in the abundance and activity of megabenthos between the two periods of study (1991–1994 and 1997–2000). In 1991–1994, the larger megabenthos occurred at an abundance of c. 71.6/ha and were dominated by large holothurians. In addition, there were very substantial populations of smaller megabenthic ophiuroids (c. 4979/ha). Together, the total megabenthos are estimated to track over some 17 cm2/m2/d (exploiting 100% of the surface of the seabed in c. 2.5 years). In 1997–2000, the larger megabenthos increased to an abundance of c. 204/ha and were joined by exceptional numbers of a small holothurian species (Amperima rosea, 6457/ha) and ophiuroids (principally Ophiocten hastatum, 53,539/ha). The total megabenthos population was tracking at an estimnated rate of c. 247 cm2/m2/d (exploiting 100% of seabed in just 6 weeks). Coincident with these increases in the abundance and activity of the megabenthos, there were apparently no mass depositions of aggregated phytodetritus to the seabed in the summers of 1997–1999. Mass occurrences of phytodetritus had been noted during the summer months of the three years previously studied (1991, 1993 and 1994), with covering between 50 and 96% of the sediment surface. There is a statistically significant (p<0.02) negative correlation between maximum extent of this seabed cover of phytodetritus and seabed tracking by megabenthos. Additional studies [Lampitt et al., Progr. Ocean. 50 (2001)], indicate that there were no substantial changes in surface ocean primary productivity, in export flux, or in the composition of the flux that might otherwise account for the apparent absence of observable concentrations of phytodetritus during the summers of 1997–1999. We postulate that the marked increase in megabenthic tracking activity resulted in the removal (via consumption, disaggregation, burial etc.) of the bulk of the incoming phytodetrital flux during these years. A simple conceptual model, based on the apparent phytodetrital fluxes observed in 1991 and 1993, suggests that the megabenthos tracking rates estimated for 1997–1999 are sufficient to account for near-total removal of this flux. However, we are not able to estimate other processes removing phytodetritus (i.e. other elements of the benthos) that may also have increased between 1991–1994 and 1997–1999. Other independent studies [e.g. Ginger et al., Progr. Ocean. 50 (2001)] of flux constituents support the possibility that just a few species of megabenthos (e.g. A. rosea, and O. hastatum) could well have consumed a major proportion of the incoming flux and so substantially modified the composition of the organic matter available to other components of the benthos.

The IOSDL DEEPSEAS programme: introduction and photographic evidence for the presence and absence of a seasonal input of phytodetritus at contrasting abyssal sites in the northeastern atlantic

Abstract This paper introduces the IOSDL DEEPSEAS programme. Two abyssal sites in the northeast Atlantic with presumed contrasting regimes of organic carbon supply have been studied. One of these sites, on the Porcupine Abyssal Plain (PAP), has an overlying water column with a winter mixed layer in excess of 500 m and was forecast to receive a highly seasonal organic input, a significant portion arriving in the form of rapidly sinking phytodetritus derived from the spring bloom. The winter mixed layer over the second site, on the Madeira Abyssal Plain (MAP), is much shallower, and the resulting flux to the benthos was expected to be quantitatively less and not in the form of aggregated phytodetritus. Recently published sediment-trap results from nearby localities indicate relatively similar total fluxes and widespread seasonality at depth, contrary to our expectations. However, benthic photographic data from the two stations seem to support the original hypothesis, at least in part. Transect photographs (and multiple-corer samples) at the PAP site in August 1989 and May 1991 revealed the presence of phytodetritus on the seafloor, relatively flocculent and evenly distributed in May and more granular and patchily distributed in August. Time-lapse photographs obtained between May 1991 and April 1992 recorded the sudden arrival of phytodetritus on 16 May and a further deposition at the beginning of June. In contrast, at the MAP site neither transect photographs in August 1990 nor time-lapse photographs obtained between August 1990 and July 1991 show evidence of the arrival of aggregated phytodetritus.

UK Atlantic Margin Environmental Survey: Introduction and overview of bathyal benthic ecology

The recent expansion of the Oil and Gas Industry in to the deep waters of the UK Atlantic Frontier prompted the industry and its regulator to reappraise the needs and means of environmental monitoring. In concert, deep-sea academics, specialist contractors, the regulator and the Industry, through the Atlantic Frontier Environmental Network (AFEN), devised and implemented a large-scale environmental survey of the deep waters to the north and west of Scotland. The AFEN-funded survey was carried out during the summers of 1996 and 1998, and involved two steps; an initial sidescan sonar mapping of the survey areas, followed up with direct seabed investigations by coring and photography. This contribution deals with the latter step. Seabed samples were collected to assess sediment type, organic content, heavy metals, hydrocarbons and macrobenthos. Photographic and video observations were employed to provide both ‘routine’ seabed assessments and to investigate particular sidescan features of note. Although essentially intended as a ‘baseline’ environmental survey, anthropogenic impacts are already evident throughout the areas surveyed. Indications of the effects of deep-sea trawling were frequently encountered (seabed trawl marks and areas of disturbed sediments), being present in almost all of the areas studied and extending to water depths in excess of 1000 m. Evidence of localised contamination of the seabed by drilling muds was also detected, though background hydrocarbon contamination is predominantly of terrestrial origin or derived from shipping. The benthic ecology of the UK Atlantic Margin is dominated by the marked differences in the hydrography of the Faroe–Shetland Channel (FSC) and the Rockall Trough (RT). Comparatively warm North Atlantic Water is common to both areas; however, in the FSC, cold (subzero) waters occupy the deeper parts of the channel (>600 m). The extreme thermal gradient present on the West Shetland Slope has a substantial influence on the distribution and diversity of the macrobenthos. While there is continuous variation in the fauna with depth, warm and cold water faunas are nonetheless quite distinct. The boundary region, centred on 400 m water depth, may be best characterised as an ecotone, having a mixed warm and cold water fauna with a distinctly enhanced diversity. The Wyville–Thomson Ridge largely prevents the cold waters of the deep FSC from entering the RT (they certainly do not influence the areas of the Malin/Hebrides Slope assessed during the survey). Consequently, the deep-water faunas north and south of the ridge are highly distinct. There is also a very marked difference in the diversity of the two faunas: diversity declines with depth in the FSC but increases with depth in the RT. The distribution of macrobenthos in the RT is largely continuous with depth, with little indication of local variations but some evidence of enhanced rates of change at around 1200 m, possibly associated with the presence of Labrador Sea Water. Other observations made during the course of the survey include: (a) the occurrence of sponge dominated communities (‘ostebund’) at mid-slope depths (ca. 500 m) north and west of Shetland; (b) the discovery of a population of sediment surface dwelling enteropneusts associated with a sandy contourite deposit at the base of the West Shetland Slope (ca. 900 m); (c) the widespread and abundant occurrence of phytodetritus in the RT but not the FSC; and (d) the discovery of the ‘Darwin Mounds’ at ca. 1000 m in the northern RT, a field of numerous, small seabed mounds that support significant growths of the coral Lophelia pertusa. These mounds also have ‘acoustically visible tails’ with dense populations of xenophyophores (Syringammina fragilissima), a species found to be common elsewhere in the RT.

Role of cold-water Lophelia pertusa coral reefs as fish habitat in the NE Atlantic

The rate of discovery of reefs of the cold-water coral Lophelia pertusa (Linnaeus, 1758) has been remarkable, and attributable to the increased use of underwater video. These reefs form a major three-dimensional habitat in deeper waters where little other ‘cover’ for fish is available. They are common in the eastern North Atlantic, and occur at least in the western North Atlantic and off central Africa. There are also other non-reef records of Lophelia in the Atlantic, and in Indian and Pacific oceans. Thus, not only are these reefs a significant habitat on a local scale, but they may also provide an important habitat over a very wide geographic scale.

The origin of deep-water, coral-topped mounds in the northern Rockall Trough, northeast Atlantic

Mounds associated with the cold water coral Lophelia pertusa are widespread in the North Atlantic, although the factors controlling their distribution are not well understood. Here we examine a group of small, coral-topped mounds (the Darwin mounds) which occur at 1000 m water depth in the northern Rockall Trough, northwest of the UK. Individual mounds are up to 75 m in diameter and 5 m high, although some ‘mound-like’ targets seen on sidescan sonar have little or even negative relief. Some mounds are associated with ‘tail-like’ features, imaged as elongate patches of moderate backscatter up to 500 m long, elongated parallel to prevailing bottom currents. High-resolution sidescan images and seabed photographs show hundreds of coral colonies, each a metre or so across, on each individual mound. Many other organisms, mainly suspension feeders, occur in association with the coral. Piston cores from the mounds contain predominantly quartz sand with only scattered coral fragments, showing that bioclastic material is not a major contributor to mound building. A field of seabed pockmarks occurs immediately south of the Darwin mounds. On sidescan sonar data, pockmarks are low relief, circular depressions, typically around 50 m in diameter. The seafloor around the pockmarks consists of uniform, heavily-burrowed, muddy sediments and no specific biological communities, nor any sedimentological or photographic evidence for active seepage, were observed. The distribution of mounds and pockmarks suggests a gradual transition from mounds in the north to pockmarks in the south. This, combined with the lack of bioclastic material in the mound sediments, suggests that both mounds and pockmarks are created by fluid escape from below the seafloor. Mounds occur where fluids carry subsurface sand to the surface, where it forms mounds because bottom currents are not strong enough to disperse it. Pockmarks form where muddy material is eroded by fluid escape but dispersed by bottom currents. Despite the origin of mounds through fluid escape, we suggest that it is the elevated mound topography, rather than any fluid escape, that is advantageous to the corals. This is supported: (1) by the wide variety of suspension-feeding organisms that occur on the mounds, since all of these are unlikely to have a specialised seepage-related lifestyle, and (2) because corals and their associated community do not occur around pockmarks, where seepage has also occurred but elevated topography is absent.