Subtle variation in abyssal terrain induces significant change in benthic megafaunal abundance, diversity, and community structure

Bathymetric gradients in the deep sea are known to affect key benthic community characteristics such as diversity. However, most studies investigate large-scale bathymetric variation, while habitat heterogeneity related to modest bathymetric variation has generally been overlooked because of limitations to sampling technology. We investigate the role of modest bathymetric variation (~10 m water depth intervals) on an abyssal hill, and horizontal variation at the 0.1–10 km scale, in the structuring of abyssal megafaunal assemblages. We assess numerical density, biomass density, diversity, and assemblage composition using seabed photographs captured with an autonomous underwater vehicle and sediment characteristics determined from cores. We detect significant differences in sediment particle size and organic carbon content, in relation to modest topographic elevation, with a greater fraction of fine particles and organic carbon on the abyssal plain than the hill. Total megafaunal numerical and biomass density, diversity, and the numerical densities of feeding groups were significantly different with modest topographic elevation; similarly, megafaunal composition varied significantly between ~10 m depth intervals. In relation to mesoscale horizontal variation, we also record significant differences between megabenthic communities in two abyssal plain areas with no significant differences in measured sedimentary characteristics and only a 2 m difference in water depth. Differences in these communities were detected in terms of dominance, assemblage composition by density and biomass, and numerical densities of feeding groups. These observations strongly indicate that previous general concepts of the abyssal environment greatly underestimate this mesoscale heterogeneity, such that beta- and gamma-diversity in the abyss may be higher than estimated. Importantly, these results also have clear implications for the design and interpretation of environmental survey and monitoring programmes in the abyss

The hemisessile lifestyle and feeding strategies of Iosactis vagabunda (Actiniaria, Iosactiidae), a dominant megafaunal species of the Porcupine Abyssal Plain

Iosactis vagabunda Riemann-Zürneck, 1997 (Actiniaria, Iosactiidae) is a small endomyarian anemone, recently quantified as the greatest contributor to megafaunal density (48%; 2372 individuals ha−1) on the Porcupine Abyssal Plain (PAP). We used time-lapse photography to observe 18 individuals over a period of approximately 20 months at 8-h intervals, and one individual over 2 weeks at 20-mine intervals, and report observations on its burrowing activity, and both deposit and predatory feeding behaviours. We recorded the apparent subsurface movement of an individual from an abandoned burrow to a new location, and burrow creation there. Raptorial deposit feeding on settled phytodetritus particles was observed, as was predation on a polychaete 6-times the estimated biomass of the anemone. Though essentially unnoticed in prior studies of the PAP, I. vagabunda may be a key component of the benthic community, and may make a critical contribution to the carbon cycling at the PAP long-term time-series study site

Abundance and morphology of Paleodictyon nodosum, observed at the Clarion-Clipperton Zone

Paleodictyon is an important trace fossil characterised by a regular hexagonal structure and typical of ancient deep-ocean habitats as far back as the Ordovician. It is represented in modern deep-sea settings by Paleodictyon nodosum, known from the Mid-Atlantic Ridge, the South Atlantic, and off eastern Australia. Here we report the occurrence of P. nodosum in the Clarion Clipperton Zone (CCZ), abyssal equatorial Pacific, an area characterised by polymetallic nodule fields. At the study site within the International Seabed Authority northeastern Area of Particular Environmental Interest (APEI-6), P. nodosum appeared as a compact, regular pattern of small circular openings on the seafloor, each pattern interpreted as reflecting the activity of an individual organism. The patterns had a mean size (maximum dimension) of 45 mm?±?16 mm SD (n?=?841) and occurred at a density of 0.33 individuals m?2. Most (82%) were interrupted by nodules, but those that were not displayed both regular (59%) and irregular (41%) forms, the former having equal numbers of rows along the three axes (6 x 6 x 6 and 8 x 8 x 8). In both size and morphology, our Paleodictyon traces were more similar to the Australian examples than to those from the Mid-Atlantic Ridge

Xenophyophores (Rhizaria, Foraminifera), including four new species and two new genera, from the western Clarion-Clipperton Zone (abyssal equatorial Pacific)

The Clarion-Clipperton Zone (CCZ) occupies a vast swathe of the Pacific with extensive polymetallic nodule deposits. Eastern and central parts host diverse assemblages of xenophyophores (megafaunal agglutinated foraminifera). Here we describe xenophyophores obtained using a Remotely Operated Vehicle from the western CCZ. Eleven distinct forms include two known species, Stannophyllum zonarium Haeckel, 1888 and Aschemonella monile Gooday and Holzmann in Gooday et al., 2017b. Another four are described as new species based on morphological and genetic data. In Abyssalia foliformis gen. nov., sp. nov. and Abyssalia sphaerica sp. nov. the flattened or spherical test comprises a homogeneous framework of sponge spicules. Psammina tenuis sp. nov. has a delicate, thin, plate-like test. Moanammina semicircularis gen. nov., sp. nov. has a stalked, fan-shaped test and is genetically identical to ‘Galatheammina sp. 6’ of Gooday and co-workers from the eastern CCZ. Sequence data revealed a spherical ‘mudball’, which disintegrated and cannot be formally described, to be a novel xenophyophore. Finally, four morphospecies are represented by dead tests: Psammina spp., Reticulammina sp., and an unknown genus with a unique test structure. This collection enhances our knowledge of Pacific xenophyophore diversity and provides the first genetic confirmation of wide geographic ranges for abyssal species

Abyssal hills - hidden source of increased habitat heterogeneity, benthic megafaunal biomass and diversity in the deep sea

Abyssal hills are the most abundant landform on Earth, yet the ecological impact of the resulting habitat heterogeneity on the wider abyss is largely unexplored. Topographic features are known to influence food availability and the sedimentary environment in other deep-sea habitats, in turn affecting the species assemblage and biomass. To assess this spatial variation, benthic assemblages and environmental conditions were compared at four hill and four plain sites at the Porcupine Abyssal Plain. Here we show that differences in megabenthic communities on abyssal hills and the adjacent plain are related to environmental conditions, which may be caused by local topography and hydrodynamics. Although these hills may receive similar particulate organic carbon flux (food supply from the surface ocean) to the adjacent plain, they differ significantly in depth, slope, and sediment particle size distribution. We found that megafaunal biomass was significantly greater on the hills (mean 13.45 g m−2, 95% confidence interval 9.25–19.36 g m−2) than the plain (4.34 g m−2, 95% CI 2.08–8.27 g m−2; ANOVA F(1, 6) = 23.8, p < 0.01). Assemblage and trophic compositions by both density and biomass measures were significantly different between the hill and plain, and correlated with sediment particle size distributions. Hydrodynamic conditions responsible for the local sedimentary environment may be the mechanism driving these assemblage differences. Since the ecological heterogeneity provided by hills in the abyss has been underappreciated, regional assessments of abyssal biological heterogeneity and diversity may be considerably higher than previously thought

Environmental Impact Assessment process for deep-sea mining in ‘the Area’

Environmental Impact Assessment (EIA) is key to the robust environmental management of industrial projects; it is used to anticipate, assess and reduce environmental and social risks of a project. It is instrumental in project planning and execution, and often required for financing and regulatory approval to be granted. The International Seabed Authority currently requires an EIA for deep-sea mining (DSM) in areas beyond national jurisdiction (the Area), but the existing regulations present only a portion of a robust EIA process. This article presents an ideal EIA process for DSM, drawing upon the application of EIA from allied industries. It contains screening, scoping and assessment phases, along with the development of an environmental management plan. It also includes external review by experts, stakeholder consultation, and regulatory review. Lessons learned from application of EIA elsewhere are discussed in relation to DSM, including the integration of EIA into UK domestic law, and the reception of EIAs prepared for seabed ore extraction in the Exclusive Economic Zones of New Zealand and Papua New Guinea. Finally, four main challenges of implementing the EIA process to DSM in the Area are presented: 1) EIA process for DSM needs to incorporate mechanisms to address uncertainty; 2) detailed requirements for the EIA process phases should be made clear; 3) mechanisms are needed to ensure that the EIA influences decision making; and, 4) the EIA process requires substantial input and involvement from the regulator

Giant, highly diverse protists in the abyssal Pacific: vulnerability to impacts from seabed mining and potential for recovery

Xenophyophores, giant deep-sea agglutinated foraminifera, dominate the benthic megafauna in the eastern equatorial Pacific Clarion-Clipperton Zone. This abyssal (>4000 m depth) region hosts major deposits of polymetallic nodules targeted for future seabed mining, an activity that would destroy these highly diverse and delicate protists, particularly those living on the nodules themselves. Since the cell occupies only a small proportion of their test volume, xenophyophores may make a fairly modest contribution to benthic biomass and carbon cycling. Nevertheless, xenophyophore tests can passively enhance particle deposition, concentrate food, and provide habitat structure utilized by diverse organisms. Their destruction could therefore influence the recovery of benthic communities. Species requiring nodule substrates will likely not recover, since nodules take millions of years to form. However, xenophyophores can grow quickly and colonize extensive volcanic ash deposits within years, suggesting that sediment-dwelling species could be among the first large immobile organisms to reappear in mining-impacted areas

Response of deep-sea deposit-feeders to detrital inputs: A comparison of two abyssal time-series sites

Biological communities on the abyssal plain are largely dependent on detritus from the surface ocean as their main source of energy. Seasonal fluctuations in the deposition of that detritus cause temporal variations in the quantity and quality of food available to these communities, altering their structure and the activity of the taxa present. However, direct observations of energy acquisition in relation to detritus availability across megafaunal taxa in abyssal communities are few. We used time-lapse photography and coincident measurement of organic matter flux from water column sediment traps to examine the impact of seasonal detrital inputs on resource acquisition by the deposit feeding megafauna assemblages at two sites: Station M (Northeast Pacific, 4000 m water depth) and the Porcupine Abyssal Plain Sustained Observatory (PAP-SO, Northeast Atlantic 4850 m water depth). At Station M, studied over 18-months, the seasonal particle flux was followed by a salp deposition event. At that site, diversity in types of deposit feeding was related to seabed cover by detritus. At PAP-SO, studied over 30 months, the seasonal particle flux consisted of two peaks annually. While the two study sites were similar in mean flux (~8.0 mgC m−2 d−1), the seasonality in the flux was greater at PAP-SO. The mean overall tracking at PAP-SO was five times that of Station M (1.9 and 0.4 cm2 h−1, respectively); both are likely underestimated because tracking by some common taxa at both sites could not be quantified. At both sites, responses of deposit-feeding megafauna to the input of detritus were not consistent across the taxa studied. The numerically-dominant megafauna (e.g. echinoids, large holothurians and asteroids) did not alter their deposit feeding in relation to the seasonality in detrital supply. Taxa for which deposit feeding occurrence or rate were correlated to seasonality in particle flux were relatively uncommon (e.g. enteropneusta), known to cache food (e.g. echiurans), or to be highly selective for fresh detritus (e.g. the holothurian Oneirophanta mutabilis). Thus, the degree of seasonality in deposit feeding appeared to be taxon-specific and related to natural history characteristics such as feeding and foraging modes

Environmental considerations for impact and preservation reference zones for deep-sea polymetallic nodule mining

Development of guidance for environmental management of the deep-sea mining industry is important as contractors plan to move from exploration to exploitation activities. Two priorities for environmental management are monitoring and mitigating the impacts and effects of activities. International regulation of deep-sea mining activities stipulates the creation of two types of zones for local monitoring within a claim, impact reference zones (IRZ) and preservation reference zones (PRZ). The approach used for allocating and assessing these zones will affect what impacts can be measured, and hence taken into account and managed. This paper recommends key considerations for establishing these reference zones for polymetallic nodule mining. We recommend that zones should be suitably large (Recommendation 1) and have sufficient separation (R2) to allow for repeat monitoring of representative impacted and control sites. Zones should be objectively defined following best-practice and statistically robust approaches (R3). This will include the designation of multiple PRZ and IRZ (R4) for each claim. PRZs should be representative of the mined area, and thus should contain high -quality resource (R5) but PRZs in other habitats could also be valuable (R6). Sediment plumes will influence design of PRZ and may need additional IRZ to monitor their effects (R7), which may extend beyond the boundaries of a claim (R8). The impacts of other expected changes should be taken into account (R9). Sharing PRZ design, placement, and monitoring could be considered amongst adjacent claims (R10). Monitoring should be independently verified to enhance public trust and stakeholder support (R11)

First in-situ monitoring of sponge response and recovery to an industrial sedimentation event

Assessment of risks to seabed habitats from industrial activities is based on the resilience and potential for recovery. Increased sedimentation, a key impact of many offshore industries, results in burial and smothering of benthic organisms. Sponges are particularly vulnerable to increases in suspended and deposited sediment, but response and recovery have not been observed in-situ. We quantified the impact of sedimentation from offshore hydrocarbon drilling over ∼5 days on a lamellate demosponge, and its recovery in-situ over ∼40 days using hourly time-lapse photographs with measurements of backscatter (a proxy of suspended sediment) and current speed. Sediment accumulated on the sponge then cleared largely gradually but occasionally sharply, though it did not return to the initial state. This partial recovery likely involved a combination of active and passive removal. We discuss the use of in-situ observing, which is critical to monitoring impacts in remote habitats, and need for calibration to laboratory conditions