Ashley A. Rowden

Man and the Last Great Wilderness: Human Impact on the Deep Sea

The deep sea, the largest ecosystem on Earth and one of the least studied, harbours high biodiversity and provides a wealth of resources. Although humans have used the oceans for millennia, technological developments now allow exploitation of fisheries resources, hydrocarbons and minerals below 2000 m depth. The remoteness of the deep seafloor has promoted the disposal of residues and litter. Ocean acidification and climate change now bring a new dimension of global effects. Thus the challenges facing the deep sea are large and accelerating, providing a new imperative for the science community, industry and national and international organizations to work together to develop successful exploitation management and conservation of the deep-sea ecosystem. This paper provides scientific expert judgement and a semi-quantitative analysis of past, present and future impacts of human-related activities on global deep-sea habitats within three categories: disposal, exploitation and climate change. The analysis is the result of a Census of Marine Life – SYNDEEP workshop (September 2008). A detailed review of known impacts and their effects is provided. The analysis shows how, in recent decades, the most significant anthropogenic activities that affect the deep sea have evolved from mainly disposal (past) to exploitation (present). We predict that from now and into the future, increases in atmospheric CO2 and facets and consequences of climate change will have the most impact on deep-sea habitats and their fauna. Synergies between different anthropogenic pressures and associated effects are discussed, indicating that most synergies are related to increased atmospheric CO2 and climate change effects. We identify deep-sea ecosystems we believe are at higher risk from human impacts in the near future: benthic communities on sedimentary upper slopes, cold-water corals, canyon benthic communities and seamount pelagic and benthic communities. We finalise this review with a short discussion on protection and management methods.

Submarine canyons: hotspots of benthic biomass and productivity in the deep sea.

Submarine canyons are dramatic and widespread topographic features crossing continental and island margins in all oceans. Canyons can be sites of enhanced organic-matter flux and deposition through entrainment of coastal detrital export, dense shelf-water cascade, channelling of resuspended particulate material and focusing of sediment deposition. Despite their unusual ecological characteristics and global distribution along oceanic continental margins, only scattered information is available about the influence of submarine canyons on deep-sea ecosystem structure and productivity. Here, we show that deep-sea canyons such as the Kaikoura Canyon on the eastern New Zealand margin (42°01′ S, 173°03′ E) can sustain enormous biomasses of infaunal megabenthic invertebrates over large areas. Our reported biomass values are 100-fold higher than those previously reported for deep-sea (non-chemosynthetic) habitats below 500 m in the ocean. We also present evidence from deep-sea-towed camera images that areas in the canyon that have the extraordinary benthic biomass also harbour high abundances of macrourid (rattail) fishes likely to be feeding on the macro- and megabenthos. Bottom-trawl catch data also indicate that the Kaikoura Canyon has dramatically higher abundances of benthic-feeding fishes than adjacent slopes. Our results demonstrate that the Kaikoura Canyon is one of the most productive habitats described so far in the deep sea. A new global inventory suggests there are at least 660 submarine canyons worldwide, approximately 100 of which could be biomass hotspots similar to the Kaikoura Canyon. The importance of such deep-sea canyons as potential hotspots of production and commercial fisheries yields merits substantial further study.

Static and dynamic environmental factors determining the community structure of estuarine macrobenthos in SW Britain: why is the Severn estuary different?

(1) A survey of intertidal macrobenthic invertebrates at forty sites in six estuaries in SW Britain was undertaken to identify the key environmental variables which determine the composition of these communities and to provide a prediction of conditions in the Severn estuary should a barrage for the generation of tidal power be built. (2) Multi-dimensional Scaling Ordination of the faunistic data showed that the sites were separated along two principal axes, one determined by static variables (e.g. sediment grain size and organic content) and the other by dynamic variables (i.e. current velocities as indexed by tidal range and wave climate as indexed by the wind fetch distance). (3) The faunistic composition at sites in the Severn estuary is significantly different from the other five estuaries, which by and large are not significantly different from each other. (4) The environmental variables responsible for the uniqueness of the Severn are dynamic rather than static, and should be predictable precisely in the post-barrage Severn. (5) Implications for qualitative changes in the fauna are discussed.

Effect of habitat fragmentation on the macroinvertebrate infaunal communities associated with the seagrass Zostera marina L.

1. The effect of habitat fragmentation was investigated in two adjacent, yet separate, intertidal Zostera marina beds in the Salcombe Estuary, Devon, UK. The seagrass bed on the west bank comprised a continuous meadow of ca. 2.3 ha, whilst the bed on the east bank of the estuary was fragmented into patches of 6–9 m2. 2. Three 10 cm diameter core samples for infaunal macroinvertebrates were taken from three stations within each bed. No significant difference was found in univariate community parameters between beds, or in measured seagrass parameters. However, multivariate analysis revealed a significant difference in community composition, due mainly to small changes in species abundance rather than differences in the species present. 3. The species contributing most to the dissimilarity between the two communities were polychaetes generally associated with unvegetated habitats (e.g. Magelona mirabilis) and found to be more common in the fragmented bed. 4. A significant difference in median grain size and sorting coefficient was recorded between the two beds, and median grain size was found to be the variable best explaining multivariate community patterns. 5. The results of the study provide evidence for the effects of habitat fragmentation on the communities associated with seagrass beds, habitats which are of high conservation importance. As the infaunal community is perhaps intuitively the component least likely to be affected by fragmentation at the scale observed, the significant difference in community composition recorded has consequences for more sensitive and high-profile parts of the biota (e.g. fish), and thus for the conservation of seagrass habitats and their associated communities. Copyright

Physical characterisation and a biologically focused classification of “seamounts” in the New Zealand region

Abstract The physical, biological, and oceano‐graphic characteristics of seamounts of the New Zealand region of the South Pacific Ocean are poorly known. The aim of this study was to present a synopsis of the physical characteristics of seamounts within the region, and to present a preliminary classification using biologically meaningful variables. Data for up to 16 environmental variables were collated and used to describe the distribution and characteristics of the c. 800 known seamounts in the New Zealand region. Seamounts span a wide range of sizes, depths, elevation, geological associations and origins, and occur over the latitudinal range of the region, lying in different water masses of varying productivity, and both near shore and off shore. As such, it was difficult to generally describe New Zealand seamounts, as there is no “typical” feature. Thirteen environmental variables were included in a multivariate cluster analysis to identify 12 seamount similarity groupings, for a subset of over half the known seamounts. The groupings generally displayed an appreciable geographic distribution throughout the region, and were largely characterised by a combination of four variables (depth at peak, depth at base, elevation, and distance from continental shelf). In the future, the findings of the present study can be tested to determine the validity and usefulness of the approach for directing future biodiversity research and informing management of seamount habitat.

Marine fish may be biochemically constrained from inhabiting the deepest ocean depths

Significance Fish appear to be absent from the oceans greatest depths, the trenches from 8,400–11,000 m. The reason is unknown, but hydrostatic pressure is suspected. We propose that the answer is the need for high levels of trimethylamine oxide (TMAO, common in many marine animals), a potent stabilizer capable of counteracting the destabilization of proteins by pressure. TMAO is known to increase with depth in bony fishes (teleosts) down to 4,900 m. By capturing the worlds second-deepest known fish, the hadal snailfish Notoliparis kermadecensis from 7,000 m, we find that they have the highest recorded TMAO contents, which, moreover, yield an extrapolated maximum for fish at about 8,200 m. This is previously unidentified evidence that biochemistry may constrain depth for a large taxonomic group. No fish have been found in the deepest 25% of the ocean (8,400–11,000 m). This apparent absence has been attributed to hydrostatic pressure, although direct evidence is wanting because of the lack of deepest-living species to study. The common osmolyte trimethylamine N-oxide (TMAO) stabilizes proteins against pressure and increases with depth, going from 40 to 261 mmol/kg in teleost fishes from 0 to 4,850 m. TMAO accumulation with depth results in increasing internal osmolality (typically 350 mOsmol/kg in shallow species compared with seawaters 1,100 mOsmol/kg). Preliminary extrapolation of osmolalities of predicted isosmotic state at 8,000–8,500 m may indicate a possible physiological limit, as greater depths would require reversal of osmotic gradients and, thus, osmoregulatory systems. We tested this prediction by capturing five of the second-deepest known fish, the hadal snailfish (Notoliparis kermadecensis; Liparidae), from 7,000 m in the Kermadec Trench. We found their muscles to have a TMAO content of 386 ± 18 mmol/kg and osmolality of 991 ± 22 mOsmol/kg. These data fit previous extrapolations and, combined with new osmolalities from bathyal and abyssal fishes, predict isosmotic state at 8,200 m. This is previously unidentified evidence that biochemistry could constrain the depth of a large, complex taxonomic group.

Science priorities for seamounts: research links to conservation and management.

Seamounts shape the topography of all ocean basins and can be hotspots of biological activity in the deep sea. The Census of Marine Life on Seamounts (CenSeam) was a field program that examined seamounts as part of the global Census of Marine Life (CoML) initiative from 2005 to 2010. CenSeam progressed seamount science by collating historical data, collecting new data, undertaking regional and global analyses of seamount biodiversity, mapping species and habitat distributions, challenging established paradigms of seamount ecology, developing new hypotheses, and documenting the impacts of human activities on seamounts. However, because of the large number of seamounts globally, much about the structure, function and connectivity of seamount ecosystems remains unexplored and unknown. Continual, and potentially increasing, threats to seamount resources from fishing and seabed mining are creating a pressing demand for research to inform conservation and management strategies. To meet this need, intensive science effort in the following areas will be needed: 1) Improved physical and biological data; of particular importance is information on seamount location, physical characteristics (e.g. habitat heterogeneity and complexity), more complete and intensive biodiversity inventories, and increased understanding of seamount connectivity and faunal dispersal; 2) New human impact data; these shall encompass better studies on the effects of human activities on seamount ecosystems, as well as monitoring long-term changes in seamount assemblages following impacts (e.g. recovery); 3) Global data repositories; there is a pressing need for more comprehensive fisheries catch and effort data, especially on the high seas, and compilation or maintenance of geological and biodiversity databases that underpin regional and global analyses; 4) Application of support tools in a data-poor environment; conservation and management will have to increasingly rely on predictive modelling techniques, critical evaluation of environmental surrogates as faunal “proxies”, and ecological risk assessment.

Critical-Evaluation Of Sediment Turnover Estimates For Callianassidae (Decapoda Thalassinidea)

Abstract Members of the decapod family Callianassidae influence sediment dynamics and ecosystem function via their bioturbation activities. The latter is currently assessed by measures of sediment turnover rates which are collected, calculated and expressed by different methods. Some estimates, particularly extrapolations which do not consider the influence of temperature, population structure and expulsion behaviour, are likely to be significantly over/under estimates of sediment turnover. Therefore, published values of sediment turnover by the Callianassidae need to be treated with caution and are not strictly comparable. In view of the need for assessing the relative importance of callianassid bioturbation, it is suggested that attempts should be made to standardize the measurement of sediment turnover rate to allow future comparisons to be made with confidence.

Factors influencing sediment turnover by the burrowing ghost shrimp Callianassa filholi (Decapoda : Thalassinidea)

Abstract Bioturbation by the burrowing thalassinidean shrimp Callianassa filholi (Milne-Edwards 1878) was studied at an intertidal sandflat in Otago Harbour, south-eastern New Zealand, over a period of 12 months. The amount of sediment expelled from shrimp burrows was measured each month (by direct entrapment over 24 h) and inhabiting shrimp were subsequently captured from the burrows. Rate of sediment expulsion was significantly related to seawater temperature, shore height of burrow and time, whilst the amount of sediment expelled from individual burrows was positively related to size of inhabiting shrimp but independent of sex. Annual sediment turnover for C. filholi was estimated at 96 kg (dry) m −2 year −1 which accounted for the variability imposed by monthly changes in seawater temperature and the spatial and temporal dynamics of the shrimp population. The results demonstrate the need to take account of physical and biological factors when constructing sediment turnover estimates for callianassid shrimps, and inferring their bioturbatory significance.

A contribution to the biology of the burrowing mud shrimp, Callianassa subterranea (Decapoda: Thalassinidea)

Samples of the mud shrimp Callianassa subterranea (Montagu) were taken at irregular intervals (September and October 1989, April and July 1990, August 1991) from a depth of 47 m at a fixed station in the North Sea (54°35′N 04°50′E). At this site, mud shrimps appear to have a contagious distribution, the mean density varied between 38 and 59 individuals m -2 and the sex ratio was biased significantly to males (mean male: female ratio, 1·9), except for shrimps of ≥8 mm carapace length (CL) which had an equal sex ratio. The combined samples for the full sampling period showed a bimodal distribution for males and females, with a possible third mode for males, suggesting a life cycle of between 2 and 3 years. Only females ≥7 mm CL carried eggs. Lack of samples over winter prevent any conclusive description of seasonal population patterns; however, reproduction (based on the presence of ovigerous females) extended from April to September with a peak in July and post-larval shrimps ( Ione thoracica (Montagu) infected between 20% (October) and 10–8% (August) of mud shrimps, with males having a significantly higher level of infestation than females.