Discovery of widely available abyssal rock patches reveals overlooked habitat type and prompts rethinking deep-sea biodiversity.

Habitat heterogeneity and species diversity are often linked. On the deep seafloor, sediment variability and hard-substrate availability influence geographic patterns of species richness and turnover. The assumption of a generally homogeneous, sedimented abyssal seafloor is at odds with the fact that the faunal diversity in some abyssal regions exceeds that of shallow-water environments. Here we show, using a ground-truthed analysis of multibeam sonar data, that the deep seafloor may be much rockier than previously assumed. A combination of bathymetry data, ruggedness, and backscatter from a trans-Atlantic corridor along the Vema Fracture Zone, covering crustal ages from 0 to 100 Ma, show rock exposures occurring at all crustal ages. Extrapolating to the whole Atlantic, over 260,000 km2 of rock habitats potentially occur along Atlantic fracture zones alone, significantly increasing our knowledge about abyssal habitat heterogeneity. This implies that sampling campaigns need to be considerably more sophisticated than at present to capture the full deep-sea habitat heterogeneity and biodiversity.

Evolution and phylogeny of the deep-sea isopod families Desmosomatidae Sars, 1897 and Nannoniscidae Hansen, 1916 (Isopoda: Asellota).

In the deep sea, the phylogeny and biogeography of only a few taxa have been well studied. Although more than 200 species in 32 genera have been described for the asellote isopod families Desmosomatidae Sars, 1897 and Nannoniscidae Hansen, 1916 from all ocean basins, their phylogenetic relationships are not completely understood. There is little doubt about the close relationship of these families, but the taxonomic position of a number of genera is so far unknown. Based on a combined morphological phylogeny using the Hennigian method with a dataset of 107 described species and a molecular phylogeny based on three markers (COI, 16S, and 18S) with 75 species (most new to science), we could separate Desmosomatidae and Nannoniscidae as separate families. However, we could not support the concept of the subfamilies Eugerdellatinae Hessler, 1970 and Desmosomatinae Hessler, 1970. Most genera of both families were well supported, but several genera appear as para- or even polyphyletic. Within both families, convergent evolution and analogies caused difficulty in defining apomorphies for phylogenetic reconstructions and this is reflected in the results of the concatenated molecular tree. There is no biogeographic pattern in the distribution as the genera occur over the entire Atlantic and Pacific Ocean, showing no specific phylogeographical pattern. Poor resolution at deep desmosomatid nodes may reflect the long evolutionary history of the family and rapid evolutionary radiations. Supplementary Information: The online version contains supplementary material available at 10.1007/s13127-021-00509-9.

Phylogenetic and functional evidence suggests that deep-ocean ecosystems are highly sensitive to environmental change and direct human disturbance.

An understanding of the balance of interspecific competition and the physical environment in structuring organismal communities is crucial because those communities structured primarily by their physical environment typically exhibit greater sensitivity to environmental change than those structured predominantly by competitive interactions. Here, using detailed phylogenetic and functional information, we investigate this question in macrofaunal assemblages from Northwest Atlantic Ocean continental slopes, a high seas region projected to experience substantial environmental change through the current century. We demonstrate assemblages to be both phylogenetically and functionally under-dispersed, and thus conclude that the physical environment, not competition, may dominate in structuring deep-ocean communities. Further, we find temperature and bottom trawling intensity to be among the environmental factors significantly related to assemblage diversity. These results hint that deep-ocean communities are highly sensitive to their physical environment and vulnerable to environmental perturbation, including by direct disturbance through fishing, and indirectly through the changes brought about by climate change.

Dependency of Antarctic zooplankton species on ice algae-produced carbon suggests a sea ice-driven pelagic ecosystem during winter.

How the abundant pelagic life of the Southern Ocean survives winter darkness, when the sea is covered by pack ice and phytoplankton production is nearly zero, is poorly understood. Ice-associated ("sympagic") microalgae could serve as a high-quality carbon source during winter, but their significance in the food web is so far unquantified. To better understand the importance of ice algae-produced carbon for the overwintering of Antarctic organisms, we investigated fatty acid (FA) and stable isotope compositions of 10 zooplankton species, and their potential sympagic and pelagic carbon sources. FA-specific carbon stable isotope compositions were used in stable isotope mixing models to quantify the contribution of ice algae-produced carbon (αIce ) to the body carbon of each species. Mean αIce estimates ranged from 4% to 67%, with large variations between species and depending on the FA used for the modelling. Integrating the αIce estimates from all models, the sympagic amphipod Eusirus laticarpus was the most dependent on ice algal carbon (αIce : 54%-67%), and the salp Salpa thompsoni showed the least dependency on ice algal carbon (αIce : 8%-40%). Differences in αIce estimates between FAs associated with short-term vs. long-term lipid pools suggested an increasing importance of ice algal carbon for many species as the winter season progressed. In the abundant winter-active copepod Calanus propinquus, mean αIce reached more than 50% in late winter. The trophic carbon flux from ice algae into this copepod was between 3 and 5 mg C m-2  day-1 . This indicates that copepods and other ice-dependent zooplankton species transfer significant amounts of carbon from ice algae into the pelagic system, where it fuels the food web, the biological carbon pump and elemental cycling. Understanding the role of ice algae-produced carbon in these processes will be the key to predictions of the impact of future sea ice decline on Antarctic ecosystem functioning.

Commonness and rarity in the marine biosphere.

Explaining patterns of commonness and rarity is fundamental for understanding and managing biodiversity. Consequently, a key test of biodiversity theory has been how well ecological models reproduce empirical distributions of species abundances. However, ecological models with very different assumptions can predict similar species abundance distributions, whereas models with similar assumptions may generate very different predictions. This complicates inferring processes driving community structure from model fits to data. Here, we use an approximation that captures common features of "neutral" biodiversity models--which assume ecological equivalence of species--to test whether neutrality is consistent with patterns of commonness and rarity in the marine biosphere. We do this by analyzing 1,185 species abundance distributions from 14 marine ecosystems ranging from intertidal habitats to abyssal depths, and from the tropics to polar regions. Neutrality performs substantially worse than a classical nonneutral alternative: empirical data consistently show greater heterogeneity of species abundances than expected under neutrality. Poor performance of neutral theory is driven by its consistent inability to capture the dominance of the communities' most-abundant species. Previous tests showing poor performance of a neutral model for a particular system often have been followed by controversy about whether an alternative formulation of neutral theory could explain the data after all. However, our approach focuses on common features of neutral models, revealing discrepancies with a broad range of empirical abundance distributions. These findings highlight the need for biodiversity theory in which ecological differences among species, such as niche differences and demographic trade-offs, play a central role.

Climate change and Southern Ocean ecosystems I: how changes in physical habitats directly affect marine biota.

Antarctic and Southern Ocean (ASO) marine ecosystems have been changing for at least the last 30 years, including in response to increasing ocean temperatures and changes in the extent and seasonality of sea ice; the magnitude and direction of these changes differ between regions around Antarctica that could see populations of the same species changing differently in different regions. This article reviews current and expected changes in ASO physical habitats in response to climate change. It then reviews how these changes may impact the autecology of marine biota of this polar region: microbes, zooplankton, salps, Antarctic krill, fish, cephalopods, marine mammals, seabirds, and benthos. The general prognosis for ASO marine habitats is for an overall warming and freshening, strengthening of westerly winds, with a potential pole-ward movement of those winds and the frontal systems, and an increase in ocean eddy activity. Many habitat parameters will have regionally specific changes, particularly relating to sea ice characteristics and seasonal dynamics. Lower trophic levels are expected to move south as the ocean conditions in which they are currently found move pole-ward. For Antarctic krill and finfish, the latitudinal breadth of their range will depend on their tolerance of warming oceans and changes to productivity. Ocean acidification is a concern not only for calcifying organisms but also for crustaceans such as Antarctic krill; it is also likely to be the most important change in benthic habitats over the coming century. For marine mammals and birds, the expected changes primarily relate to their flexibility in moving to alternative locations for food and the energetic cost of longer or more complex foraging trips for those that are bound to breeding colonies. Few species are sufficiently well studied to make comprehensive species-specific vulnerability assessments possible. Priorities for future work are discussed.

New species of the deep-sea munnopsid genus Tytthocope (Crustacea, Isopoda, Asellota) from the South Atlantic and the Southern Ocean.

In the benthic samples collected during the deep-sea expeditions ANDEEP from the Weddell Sea and DIVA from the Argentine Basin the isopod family Munnopsidae was the most specious and numerous. Among the collected munnopsids three new species of Tytthocope Wilson & Hessler, 1981 have been discovered. Tytthocope is one of six genera of the subfamily Eurycopinae. It differs from other genera of the Eurycopinae by having a strongly reduced pereonite 7 and inflated pleotelson. Until now the genus consisted of five described species: four of them are known from the northern Atlantic and only one species, T. sulcifrons (Barnard, 1920) was described from the southern Atlantic off the South Africa coast. The five known species of Tytthocope have been recorded from depths less than 1461 m. Descriptions of three new species of Tytthocope collected in the South Atlantic and Southern Ocean at depths between 1530-4608 m, T. divae sp. nov., T. fahrbachi sp. nov. and T. longitelson sp. nov. are presented in the paper. T. sulcifrons is redescribed herein based on the type material.

Baseline Inventory of Benthic Macrofauna in German Marine Protected Areas (2020-2022) before Closure for Bottom-Contact Fishing.

The response of benthic habitats and organisms to bottom-contact fishing intensity is investigated in marine protected areas (MPAs) of the German EEZ in the North and Baltic Seas. We examined the current state of macrofauna biodiversity in 2020-2022. Comparative analysis for macrofauna (in- and epifauna) inhabiting nine Natura 2000 MPAs constitutes a baseline to assess the effects of bottom-contact fishing exclusion in the future. Aspects of spatial and temporal variability are briefly summarized and discussed. We provide a species list for each region, including 481 taxa, of which 79 were found in both regions, 183 only in the North Sea, and 219 only in the Baltic Sea. The Baltic Sea dataset surprisingly included higher numbers of taxa and revealed more Red List species. The share of major taxonomic groups (polychaetes, bivalves and amphipods) in species richness showed peculiar commonalities between the two regions. In the North Sea, multivariate analysis of community structure revealed significantly higher within-similarity and stronger separation between the considered MPAs compared to the Baltic MPAs. Salinity, temperature and sediment fractions of sand were responsible for over 60% of the variation in the North Sea macrofauna occurrence data. Salinity, mud fraction and bottom-contact fishing were the most important drivers in the Baltic Sea and, together with other considered environmental drivers, were responsible for 53% of the variation. This study identifies aspects of macrofauna occurrence that may be used to assess (causes of) future changes.

First insights into the biodiversity and biogeography of the Southern Ocean deep sea.

Shallow marine benthic communities around Antarctica show high levels of endemism, gigantism, slow growth, longevity and late maturity, as well as adaptive radiations that have generated considerable biodiversity in some taxa. The deeper parts of the Southern Ocean exhibit some unique environmental features, including a very deep continental shelf and a weakly stratified water column, and are the source for much of the deep water in the world ocean. These features suggest that deep-sea faunas around the Antarctic may be related both to adjacent shelf communities and to those in other oceans. Unlike shallow-water Antarctic benthic communities, however, little is known about life in this vast deep-sea region. Here, we report new data from recent sampling expeditions in the deep Weddell Sea and adjacent areas (748-6,348 m water depth) that reveal high levels of new biodiversity; for example, 674 isopods species, of which 585 were new to science. Bathymetric and biogeographic trends varied between taxa. In groups such as the isopods and polychaetes, slope assemblages included species that have invaded from the shelf. In other taxa, the shelf and slope assemblages were more distinct. Abyssal faunas tended to have stronger links to other oceans, particularly the Atlantic, but mainly in taxa with good dispersal capabilities, such as the Foraminifera. The isopods, ostracods and nematodes, which are poor dispersers, include many species currently known only from the Southern Ocean. Our findings challenge suggestions that deep-sea diversity is depressed in the Southern Ocean and provide a basis for exploring the evolutionary significance of the varied biogeographic patterns observed in this remote environment.

Fissarcturus walteri sp. nov. from the Ross Sea of the Southern Ocean (Isopoda, Antarcturidae).

During the Victoria-Land Transect project two specimens of a new species, Fissarcturus walteri sp. nov., were found in the Ross Sea along the Victoria Land Coast, Antarctica, during the 19th Italica expedition in February 2004. This is the second species of Fissarcturus from the Ross Sea after F. rossi Brandt, 2007. F. walteri sp. nov. is most similar to Fissarcturus granulosus (Nordenstam, 1933) from South Georgia but can be distinguished by the blunt short, anteriorly directed supraocular spines and elevations and some other characters as described herewith.

Diversity of Southern Ocean deep-sea benthos between cosmopolitism and cryptic speciation: new species from the ANDEEP expeditions.

The richness of life in parts of the earth that to us appear inhospitable and remote never fails to fascinate scientists and non-scientists alike. The largest ecosystem of the planet, the abyssal plains of the world ocean, makes up about 90 percent of the seafloor and thus nearly 78 percent of the Earth's surface, yet only a minor fraction of this huge environment has been investigated. Authors have questioned repeatedly "how many species would live on earth and in the ocean" (Mora et al. 2011; May 2011), and recent estimations predict ~ 8.7 million (+/- 1.3 million SE) eukaryotic species globally, with 2.2 million of these being marine (Mora et al. 2011). To date, 91 % of all marine species still await description. Other authors concluded that marine biodiversity is grossly underestimated (Bouchet 2006) because so far, only one-third of all species descriptions concerns marine biota (Reaka-Kudla 1997; Groombridge & Jenkins 2000, Grassle 2001, Boltovskoy et al. 2005). We know that marine live thrives even in hadal trenches (Jamieson et al. 2009), that biogeographic ranges in the deep sea are dynamic (McClain & Mincks Hardy 2010) and that the origin of the modern deep-sea fauna is ancient (Thuy et al. 2012). For example, the origin of Isopoda dates back to Permo-Triassic times (232­ 314 mya; Lins et al. 2012).

Insights into the prokaryotic communities of the abyssal-hadal benthic-boundary layer of the Kuril Kamchatka Trench.

BACKGROUND: The Kuril-Kamchatka Trench (maximum depth 9604 m), located in the NW Pacific Ocean, is among the top seven deepest hadal trenches. The work aimed to investigate the unexplored abyssal-hadal prokaryotic communities of this fascinating, but underrated environment. RESULTS: As for the bacterial communities, we found that Proteobacteria (56.1-74.5%), Bacteroidetes (6.5-19.1%), and Actinobacteria (0.9-16.1%) were the most represented bacterial phyla over all samples. Thaumarchaeota (52.9-91.1%) was the most abundant phylum in the archaeal communities. The archaeal diversity was highly represented by the ammonia-oxidizing Nitrosopumilus, and the potential hydrocarbon-degrading bacteria Acinetobacter, Zhongshania, and Colwellia were the main bacterial genera. The α-diversity analysis evidenced that both prokaryotic communities were characterized by low evenness, as indicated by the high Gini index values (> 0.9). The ß-diversity analysis (Redundancy Analysis) indicated that, as expected, the depth significantly affected the structure of the prokaryotic communities. The co-occurrence network revealed seven prokaryotic groups that covaried across the abyssal-hadal zone of the Kuril-Kamchatka Trench. Among them, the main group included the most abundant archaeal and bacterial OTUs (Nitrosopumilus OTU A2 and OTU A1; Acinetobacter OTU B1), which were ubiquitous across the trench. CONCLUSIONS: This manuscript represents the first attempt to characterize the prokaryotic communities of the KKT abyssal-hadal zone. Our results reveal that the most abundant prokaryotes harbored by the abyssal-hadal zone of Kuril-Kamchatka Trench were chemolithotrophic archaea and heterotrophic bacteria, which did not show a distinctive pattern distribution according to depth. In particular, Acinetobacter, Zhongshania, and Colwellia (potential hydrocarbon degraders) were the main bacterial genera, and Nitrosopumilus (ammonia oxidizer) was the dominant representative of the archaeal diversity.

Journey to the deep: plastic pollution in the hadal of deep-sea trenches.

The global increase of plastic production, linked with an overall plastic misuse and waste mismanagement, leads to an inevitable increase of plastic debris that ends up in our oceans. One of the major sinks of this pollution is the deep-sea floor, which is hypothesized to accumulate in its deepest points, the hadal trenches. Little is known about the magnitude of pollution in these trenches, given the remoteness of these environments, numerous factors influencing the input and sinking behavior of plastic debris from shallower environments. This study represents to the best of our knowledge the largest survey of (macro)plastic debris sampled at hadal depths, down to 9600 m. Industrial packaging and material assignable to fishing activities were the most common debris items in the Kuril Kamchatka trench, most likely deriving from long-distance transport by the Kuroshio extension current (KE) or from regional marine traffic and fishing activities. The chemical analysis by (Attenuated Total Reflection Fourier transform infrared (ATR-FTIR) spectroscopy revealed that the main polymers detected were polyethylene (PE), polypropylene (PP) and nylon. Plastic waste is reaching the depths of the trench, although some of the items were only partially broken down. This finding suggests that complete breakdown into secondary microplastics (MP) may not always occur at the sea surface or though the water column. Due to increased brittleness, plastic debris may break apart upon reaching the hadal trench floor where plastic degrading factors were thought to be, coming off. The KKT's remote location and high sedimentation rates make it a potential site for high levels of plastic pollution, potentially making it one of the world's most heavily contaminated marine areas and an oceanic plastic deposition zone.

Deep-sea benthic crustacean and annelid data from the Bering Sea.

Samples of Crustacea and Annelida (Polychaeta, Sipuncula, and Hirudinea) were collected in the Bering Sea and the northwestern Pacific Ocean during scientific cruise SO-249 BERING in 2016. Biological samples were collected from 32 locations by the team on-board RV Sonne using a chain bag dredge at depths ranging between 330-5,070 m, and preserved in 96% ethanol. Specimens were morphologically identified to the lowest taxonomic level possible using a Leica M60 stereomicroscope. The generated data here comprise taxonomic information as well as annotated bathymetric and biogeographic information from a total of 78 samples (26 Crustacea, 47 Polychaeta, 4 Sipuncula, and 1 Hirudinea). The dataset was prepared following Darwin Core Biodiversity standards for FAIR data sharing based on Ocean Biodiversity Information System (OBIS) and Global Biodiversity Facility (GBIF) guidelines. The standardised digitised data were then mobilised to both OBIS and GBIF under CC BY 4.0 licence to publicly share and adopt the data. As records of these important marine taxa from bathyal and abyssal depths are sparse, especially from the deep Bering Sea, the herein generated and digitised data aid in filling existing knowledge gaps on their diversity and distribution in that region. As part of the "Biogeography of the NW Pacific deep-sea fauna and their possible future invasions into the Arctic Ocean" (BENEFICIAL) project, this dataset thus not only increases our knowledge in re-assessing and uncovering the deep-sea diversity of these taxa, but also serves policy and management sectors by providing first-hand data for global report assessments.

Complex sublinear burrows in the deep sea may be constructed by amphipods.

Trails, burrows, and other "life traces" in sediment provide important evidence for understanding ecology-both of the maker and of other users-and behavioral information often lacking in inaccessible ecosystems, such as the deep sea or those that are already extinct. Here, we report novel sublinear rows of openings in the abyssal plains of the North Pacific, and the first plausible hypothesis for a maker of these constructions. Enigmatic serial burrows have now been recorded in the Pacific and Atlantic deep sea. Based on image and specimen evidence, we propose that these Bering Sea excavations represent amphipod burrows, while the maker of the previously known Mid-Atlantic Ridge constructions remains undetermined. We propose that maerid amphipods could create the Pacific burrows by eating-digging horizontally below the surface along a nutrient-rich layer in the sediment, making the serial openings above them as they go, for conveniently removing excavated sediment as the excavation progresses. These striking structures contribute to local biodiversity, and their maker could be considered a deep-sea ecosystem engineer.

Northernmost (Subarctic) and deepest record of Paleodictyon: paleoecological and biological implications.

Paleodictyon is one of the most iconic and widespread of trace fossils in the geological record. However, modern examples are less well known and restricted to deep-sea settings at relatively low latitudes. Here, we report the distribution of Paleodictyon at six abyssal sites near the Aleutian Trench. This study reveals for the first time the presence of Paleodictyon at Subarctic latitudes (51°-53°N) and at depths over 4500 m, although the traces were not observed at stations deeper than 5000 m suggesting that there is some bathymetric constraint for the trace maker. Two small Paleodictyon morphotypes were recognized (average mesh size of 1.81 cm), one having a central hexagonal pattern, the other being characterized by a non-hexagonal pattern. Within the study area, Paleodictyon shows no apparent correlation with local environmental parameters. Finally, based on a worldwide morphological comparison, we conclude that the new Paleodictyon specimens represent distinct ichnospecies that are associated with the relatively eutrophic conditions in this region. Their smaller size may reflect this more eutrophic setting in which sufficient food can be obtained from a smaller area in order to satisfy the energetic requirements of the tracemakers. If so, then Paleodictyon size may provide some assistance when interpreting paleoenvironmental conditions.

Human footprints at hadal depths: interlayer and intralayer comparison of sediment cores from the Kuril Kamchatka trench.

Microplastic (MP) pollution affects almost all ecosystems on Earth. Given the increasing plastic production worldwide and the durability of these polymers, concerns arise about the fate of this material in the environment. A candidate to consider as a depositional final sink of MP is the sea floor and its deepest representatives, hadal trenches, as ultimate sinks. In this study, 13 sediment samples were collected with a multiple-corer at depths between 5740 and 9450 m from the Kuril Kamchatka trench (KKT), in the Northwest (NW) Pacific Ocean. These samples were analysed for MP presence in the upper sediment layer, by slicing the first 5 cm of sediment cores into 1 cm horizontal layers. These were compared against each other and between the sampling areas, in order to achieve a detailed picture of the depositional system of the trench and small-scale perturbations such as bioturbation. The analyses revealed the presence of 215 to 1596 MP particles per kg -1 sediment (dry weight), with a polymer composition represented by 14 polymer types and the prevalence of particles smaller than 25 µm. A heterogeneous microplastic distribution through the sediment column and different microplastic concentration and polymer types among sampling stations located in different areas of the trench reflects the dynamics of this environment and the numerous forces that drive the deposition processes and the in situ recast of this pollutant at the trench floor.

Pseudidotheaarmata sp. n., a new isopod of the genus Pseudidothea (Crustacea, Malacostraca, Isopoda) from the Atlantic sector of the Southern Ocean.

BACKGROUND: In the framework of the British Antarctic Survey (BAS) Expedion JR 15005 SO-AntEco, held in February-March 2016, the South Orkney Islands seafloor was sampled in order to investigate the distribution and composition of benthic communities around the area. NEW INFORMATION: A new species of the genus Pseudidothea Ohlin, 1901 is described from the Burdwood Bank area (South Orkney Islands). It has been collected during the SO-AntEco JR15005 RRS James Clark Ross expedition under the lead of the British Antarctic Survey (BAS). The new species, Pseudidotheaarmata sp. n., is very similar to P.scutata (Stephensen, 1947); however, it is characterised by peculiar supra-ocular spines and a different tubercular pattern. The study of the species of the Pseudidothea helps to better understand the diversity of the Pseudidotheidae in the Southern Ocean.

Patterns of eukaryotic diversity from the surface to the deep-ocean sediment.

Remote deep-ocean sediment (DOS) ecosystems are among the least explored biomes on Earth. Genomic assessments of their biodiversity have failed to separate indigenous benthic organisms from sinking plankton. Here, we compare global-scale eukaryotic DNA metabarcoding datasets (18S-V9) from abyssal and lower bathyal surficial sediments and euphotic and aphotic ocean pelagic layers to distinguish plankton from benthic diversity in sediment material. Based on 1685 samples collected throughout the world ocean, we show that DOS diversity is at least threefold that in pelagic realms, with nearly two-thirds represented by abundant yet unknown eukaryotes. These benthic communities are spatially structured by ocean basins and particulate organic carbon (POC) flux from the upper ocean. Plankton DNA reaching the DOS originates from abundant species, with maximal deposition at high latitudes. Its seafloor DNA signature predicts variations in POC export from the surface and reveals previously overlooked taxa that may drive the biological carbon pump.

Abundance data of benthic peracarid crustaceans from the South Atlantic and Southern Ocean.

Peracarid data were collected in the Southern Ocean and South Atlantic Ocean. Sampling was performed during nine different expeditions on board of RRS James Clark Ross and RV Polarstern, using epibenthic sledges (EBS) at depth ranging between 160-6348 m at 109 locations. The correlation between environmental variables and peracarid abundance was investigated. Abundance data comprise a total of 128570 peracarids (52366 were amphipods, 28516 were cumaceans, 36142 isopods, 5676 mysidaceans and 5870 were tanaidaceans). The presented data are useful to investigate the composition and abundance patterns of peracarid orders at a wide depth range and spatial scale in the Southern Ocean. They can also be reused to compare their abundance with that of other taxa in broader ecological surveys.