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

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

Essential Ocean Variables for Biogeochemical Observations

Ocean biogeochemical (BGC) processes are fundamental for several key ocean ecosystem functions and services. For example, carbon dioxide uptake and storage by chemical uptake and biological fixation in combination with transport to depth remove anthropogenic carbon from the atmosphere and counteract global warming. Another important example would be nutrient regeneration from organic matter remineralization that sustains ocean productivity and, hence, also seafood supply. Although many BGC processes are rooted in surface waters where primary productivity and ocean / atmosphere exchange takes place, the deep ocean contributes strongly to the cycling and sequestration of elements. Without a consideration of organic particle attenuation in the deep pelagic and rates of remineralization and accumulation at the vast deep seafloor, the efficiency of the biological carbon pump and the recycling of nutrients – and potential effects of global change – can hardly be quantified. Considering the relevance of the deep ocean, the Deep Ocean Observing Strategy ‘DOOS’ is revising the Essential Ocean Variables (EOVs) specified by the Global Ocean Observing System (GOOS) with a deep ocean perspective. The DOOS community recognized, that most GOOS BGC EOVs are also relevant for deep ocean observations. However, the bottom of the sea requires more attention to assess its contribution to ocean BGC. ‘Seafloor labile organic matter’ and ‘seafloor respiration’ are candidates for additional EOVs beyond the GOOS EOVs ‘oxygen’ and ‘particulate matter’. Online questionnaires are prepared to facilitate an expert revision of GOOS BGC EOVs in order to make specific suggestions for modifications to the responsible GOOS panel. The revision includes the target phenomena and spatiotemporal scales, the observing platforms and networks considered relevant, and the requirements of observations. This contribution introduces the revision approach and invites experts to take part

Benthic megafauna assemblage change over three decades in the abyss: Variations from species to functional groups

Megafaunal seafloor assemblages on the Monterey Fan in the NE Pacific (Station M, 4000 m depth) were studied between 2006–2018 using remotely operated vehicles (ROVs) as part of a continuing time-series study that began in 1989. Since 2006 we identified nearly 120,000 individual animals representing over 142 morphospecies, and observed continuous changes in the megafaunal assemblage. This study, which tracked variation in observed morphospecies over a 13-year period, is one of the most detailed long-term records of megafaunal change for abyssal depths. Our investigation shows that new variations continued to emerge, reinforcing the concept that the deep-sea is dynamic over short time scales, rather than static over long periods. Some species were uncommon, but later observed in high numbers, then decreased to very low or undetectable levels (e.g. Elpidia sp. A), while others (e.g. Psychropotes longicauda) exhibited a relatively persistent presence with less fluctuation in abundance. Decreasing total echinoderm density from 2013–2018 did not correspond with the continued occurrence of large episodic POC flux events between 2016–2018. This may be attributed to the quality of food supply arriving at the seafloor and the varied ability of organisms to utilize it. Long-term tracking (30 years) of 10 specific epibenthic echinoderm species originally quantified from camera-sled images shows a pattern of assemblage structure, perhaps returning toward the composition observed in the 1990s and early 2000s. Many questions remain as to how this abyssal site and others will change with continued, and potentially increasing, anthropogenic change in the upper ocean. For example, the marine heat anomaly known as the ‘Warm Blob’ may have influenced major ecological processes at the abyssal seafloor in terms of morphospecies and functional group composition due to changes in POC flux. The degree of dynamism continues to indicate that ad hoc or short-term investigations provide a limited perspective for assessing community structure in conservation or resource exploitation impact assessment studies in the deep sea

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

Incorporating transparency into the governance of deep-seabed mining in the Area beyond national jurisdiction

In the governance of natural resources, transparency has been linked to improved accountability, as well as enforceability, compliance, sustainability, and ultimately more equitable outcomes. Here, good practices in transparency relevant to the emerging governance of deep-seabed mining in the Area beyond national jurisdiction are identified and compared with current practices of the International Seabed Authority (ISA). The analysis found six areas of good transparency practice that could improve the accountability of deep-seabed mining: i) access to information; ii) reporting; iii) quality assurance; iv) compliance information / accreditation; v) public participation; and vi) ability to review / appeal decisions. The ISA has in some instances adopted progressive practices regarding its rules, regulations, and procedures (e.g. including the precautionary approach). However, the results here show that overall the ISA will need to consider improvements in each of the six categories above, in order to reflect contemporary best transparency practices, as well as meeting historical expectations embodied in the principle of the ‘common heritage of mankind’. This would involve a revision of its rules and procedures. The ongoing review and drafting of the ISA’s deep-seabed mining exploitation regulations offers a once-in-a-generation opportunity to improve upon the current situation. Findings from this analysis are summarised in 18 recommendations, including publication of annual reports submitted by contractors, publication of annual financial statements, development of a transparency policy, compliance reporting, and dedicated access to Committee meetings

Inter-annual species-level variations in an abyssal polychaete assemblage (Sta. M, NE Pacific, 4000 m)

Understanding the dynamics of abyssal community structure and function has become increasingly important as deep-sea resource exploitation and climate change pressures are expected to ramp up. This time-series study investigates macrofaunal polychaete dynamics at a station in the North East Pacific (Sta. M; 35˚ N 123˚ W, 4000 m, 1991-2011). Infaunal polychaete species were identified and their proxy biomass and proxy energy use rate estimated. The assemblage comprised 167 species, having a composition consistent with other abyssal areas globally. Significant changes in univariate and multivariate parameters (rank abundance distribution, Simpson’s diversity index, and species and functional group composition) were detected across 1991-2011. However, no change in biomass or energy use rate was apparent through the time-series. The largest changes in the polychaete assemblage coincided with both an increase in sinking particulate organic carbon flux to the seafloor in 2007, and a 40 km relocation of the sampling location to a site 100 m shallower, preventing a conclusive assessment of which might drive the observed variation. Analyses prior to the change of sampling location showed that the polychaete assemblage composition dynamics were primary driven by food supply variation. Changes in several species were also lagged to changes in POC flux by 4 to 10 months. The polychaete fauna exhibited a significant positive relationship between total density and total energy use rate, suggesting population-level tracking of a common resource (e.g. POC flux food supply). Neither compensatory nor energetic zero-sum dynamics were detected among the polychaete assemblage, but the results suggest that the latter occur in the macrofaunal community as a whole. The results do indicate (a) potential control of species composition, and the density of individual key species, by food supply, when the time-series prior to the sampling location was analysed separately, and (b) generally sensitive detection of environmental change by species-level analysis of the abyssal polychaete assemblage

Carbon cycling in the deep eastern North Pacific benthic food web: Investigating the effect of organic carbon input

The deep ocean benthic environment plays a role in long-term carbon sequestration. Understanding carbon cycling in the deep ocean floor is critical to evaluate the impact of changing climate on the oceanic systems. Linear inverse modeling was used to quantify carbon transfer between compartments in the benthic food web at a long time-series study site in the abyssal northeastern Pacific (Station M). Linear inverse food web models were constructed for three separate years in the time-series when particulate organic carbon (POC) flux was relatively high (1990: 0.63 mean mmol C m?2 d?1), intermediate (1995: 0.24) and low (1996: 0.12). Carbon cycling in all years was dominated by the flows involved in the microbial loop; dissolved organic carbon uptake by microbes (0.80–0.95 mean mmol C m?2 d?1), microbial respiration (0.52–0.61), microbial biomass dissolution (0.09–0.18) and the dissolution of refractory detritus (0.46–0.65). Moreover, the magnitude of carbon flows involved in the microbial loop changed in relation to POC input, with a decline in contribution during the high POC influxes, such as those recently experienced at Station M. Results indicate that during high POC episodic pulses the role of faunal mediated carbon cycling would increase. Semi-labile detritus dominates benthic faunal diets and the role of labile detritus declined with increased total POC input. Linear inverse modeling represents an effective framework to analyze high-resolution time-series data and demonstrate the impact of climate change on the deep ocean carbon cycle in a coastal upwelling system

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

Improving the estimation of deep-sea megabenthos biomass: dimension to wet weight conversions for abyssal invertebrates

Deep-sea megafaunal biomass has typically been assessed by sampling with benthic sledges and trawls, but non-destructive methods, particularly photography, are becoming increasingly common. Estimation of individual wet weight in seabed photographs has been achieved using equations obtained from measured trawl-caught specimens for a limited number of taxa. However, a lack of appropriate conversion factors has limited estimation across taxa encompassing whole communities. Here we compile relationships between measured body dimensions and preserved wet weights for a comprehensive catalogue of abyssal epibenthic megafauna, using ~47,000 specimens from the Porcupine Abyssal Plain (NE Atlantic) housed in the Discovery Collections. The practical application of the method is demonstrated using an extremely large dataset of specimen measurements from seabed photographs taken in the same location. We also collate corresponding field data on fresh wet weight, to estimate the impact of fixation in formalin and preservation in industrial denatured alcohol on the apparent biomass. Taxa with substantial proportions of soft tissues lose 35 to 60% of their wet weight during preservation, while those with greater proportions of hard tissues lose 10 to 20%. Our total estimated fresh wet weight biomass of holothurians and cnidarians in the photographic survey was ~20 times the previous estimates of total invertebrate biomass based on trawl catches. This dramatic uplift in megabenthic biomass has significant implications for studies of standing stocks, community metabolism, and numerical modelling of benthic carbon flows