David A. Bowden

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.

Hydrothermal Vents and Methane Seeps: Rethinking the Sphere of Influence

Although initially viewed as oases within a barren deep ocean, hydrothermal vent and methane seep communities are now recognized to interact with surrounding ecosystems on the sea floor and in the water column, and to affect global geochemical cycles. The importance of understanding these interactions is growing as the potential rises for disturbance from oil and gas extraction, seabed mining and bottom trawling. Here we synthesize current knowledge of the nature, extent and time and space scales of vent and seep interactions with background systems. We document an expanded footprint beyond the site of local venting or seepage with respect to elemental cycling and energy flux, habitat use, trophic interactions, and connectivity. Heat and energy are released, global biogeochemical and elemental cycles are modified, and particulates are transported widely in plumes. Hard and biotic substrates produced at vents and seeps are used by “benthic background” fauna for attachment substrata, shelter, and access to food via grazing or through position in the current, while particulates and fluid fluxes modify planktonic microbial communities. Chemosynthetic production provides nutrition to a host of benthic and planktonic heterotrophic background species through multiple horizontal and vertical transfer pathways assisted by flow, gamete release, animal movements, and succession, but these pathways remain poorly known. Shared species, genera and families indicate that ecological and evolutionary connectivity exists among vents, seeps, organic falls and background communities in the deep sea; the genetic linkages with inactive vents and seeps and background assemblages however, are practically unstudied. The waning of venting or seepage activity generates major transitions in space and time that create links to surrounding ecosystems, often with identifiable ecotones or successional stages. The nature of all these interactions is dependent on water depth, as well as regional oceanography and biodiversity. Many ecosystem services are associated with the interactions and transitions between chemosynthetic and background ecosystems, for example carbon cycling and sequestration, fisheries production, and a host of non-market and cultural services. The quantification of the sphere of influence of vents and seeps could be beneficial to better management of deep-sea environments in the face of growing industrialization.

Cold seep epifaunal communities on the Hikurangi margin, New Zealand: composition, succession, and vulnerability to human activities.

Cold seep communities with distinctive chemoautotrophic fauna occur where hydrocarbon-rich fluids escape from the seabed. We describe community composition, population densities, spatial extent, and within-region variability of epifaunal communities at methane-rich cold seep sites on the Hikurangi Margin, New Zealand. Using data from towed camera transects, we match observations to information about the probable life-history characteristics of the principal fauna to develop a hypothetical succession sequence for the Hikurangi seep communities, from the onset of fluid flux to senescence. New Zealand seep communities exhibit taxa characteristic of seeps in other regions, including predominance of large siboglinid tubeworms, vesicomyid clams, and bathymodiolin mussels. Some aspects appear to be novel; however, particularly the association of dense populations of ampharetid polychaetes with high-sulphide, high-methane flux, soft-sediment microhabitats. The common occurrence of these ampharetids suggests they play a role in conditioning sulphide-rich sediments at the sediment-water interface, thus facilitating settlement of clam and tubeworm taxa which dominate space during later successional stages. The seep sites are subject to disturbance from bottom trawling at present and potentially from gas hydrate extraction in future. The likely life-history characteristics of the dominant megafauna suggest that while ampharetids, clams, and mussels exploit ephemeral resources through rapid growth and reproduction, lamellibrachid tubeworm populations may persist potentially for centuries. The potential consequences of gas hydrate extraction cannot be fully assessed until extraction methods and target localities are defined but any long-term modification of fluid flow to seep sites would have consequences for all chemoautotrophic fauna.

Diversity and distribution of deep-sea shrimps in the Ross Sea region of Antarctica.

Although decapod crustaceans are widespread in the oceans, only Natantia (shrimps) are common in the Antarctic. Because remoteness, depth and ice cover restrict sampling in the South Ocean, species distribution modelling is a useful tool for evaluating distributions. We used physical specimen and towed camera data to describe the diversity and distribution of shrimps in the Ross Sea region of Antarctica. Eight shrimp species were recorded: Chorismus antarcticus; Notocrangon antarcticus; Nematocarcinus lanceopes; Dendrobranchiata; Pasiphaea scotiae; Pasiphaea cf. ledoyeri; Petalidium sp., and a new species of Lebbeus. For the two most common species, N. antarcticus and N. lanceopes, we used maximum entropy modelling, based on records of 60 specimens and over 1130 observations across 23 sites in depths from 269 m to 3433 m, to predict distributions in relation to environmental variables. Two independent sets of environmental data layers at 0.05° and 0.5° resolution respectively, showed how spatial resolution affected the model. Chorismus antarcticus and N. antarcticus were found only on the continental shelf and upper slopes, while N. lanceopes, Lebbeus n. sp., Dendrobranchiata, Petalidium sp., Pasiphaea cf. ledoyeri, and Pasiphaea scotiae were found on the slopes, seamounts and abyssal plain. The environmental variables that contributed most to models for N. antarcticus were depth, chlorophyll-a concentration, temperature, and salinity, and for N. lanceopes were depth, ice concentration, seabed slope/rugosity, and temperature. The relative ranking, but not the composition of these variables changed in models using different spatial resolutions, and the predicted extent of suitable habitat was smaller in models using the finer-scale environmental layers. Our modelling indicated that shrimps were widespread throughout the Ross Sea region and were thus likely to play important functional role in the ecosystem, and that the spatial resolution of data needs to be considered both in the use of species distribution models.

Diversity in a Cold Hot-Spot: DNA-Barcoding Reveals Patterns of Evolution among Antarctic Demosponges (Class Demospongiae, Phylum Porifera)

Background The approximately 350 demosponge species that have been described from Antarctica represent a faunistic component distinct from that of neighboring regions. Sponges provide structure to the Antarctic benthos and refuge to other invertebrates, and can be dominant in some communities. Despite the importance of sponges in the Antarctic subtidal environment, sponge DNA barcodes are scarce but can provide insight into the evolutionary relationships of this unique biogeographic province. Methodology/Principal Findings We sequenced the standard barcoding COI region for a comprehensive selection of sponges collected during expeditions to the Ross Sea region in 2004 and 2008, and produced DNA-barcodes for 53 demosponge species covering about 60% of the species collected. The Antarctic sponge communities are phylogenetically diverse, matching the diversity of well-sampled sponge communities in the Lusitanic and Mediterranean marine provinces in the Temperate Northern Atlantic for which molecular data are readily available. Additionally, DNA-barcoding revealed levels of in situ molecular evolution comparable to those present among Caribbean sponges. DNA-barcoding using the Segregating Sites Algorithm correctly assigned approximately 54% of the barcoded species to the morphologically determined species. Conclusion/Significance A barcode library for Antarctic sponges was assembled and used to advance the systematic and evolutionary research of Antarctic sponges. We provide insights on the evolutionary forces shaping Antarcticas diverse sponge communities, and a barcode library against which future sequence data from other regions or depth strata of Antarctica can be compared. The opportunity for rapid taxonomic identification of sponge collections for ecological research is now at the horizon.

Seafloor Habitats and Benthos of a Continental Ridge: Chatham Rise, New Zealand

Publisher Summary Chatham Rise is a submarine ridge extending ∼1,500 km due east of the South Island, New Zealand. It consists of continental basement rocks (Mesozoic graywackes) overlain by Mid- to Late Cretaceous marine sedimentary rocks and Paleocene–Miocene chalks and greensands, with sporadic Pliocene–Pleistocene intraplate volcanic edifices, especially along its northern and eastern margins. Seabed habitats are impacted by bottom trawling across most parts of the rise, with major fisheries for orange roughy, oreos ( Allocytus niger and Pseudocyttus maculatus ), and alfonsino ( Beryx splendens ) on pinnacles and slope areas along the northern and eastern flanks, hoki over water depths of 500–800 m on the flanks, and scampi ( Metanephrops challengeri ), hake ( Merluccius australis ), and ling ( Genypterus blacodes ) on the crest of the rise. Before the development of deep-water fisheries in the 1970s, Chatham Rise supported very large populations of orange roughy. These populations are now considerably reduced and larger aggregations, particularly in spawning season, are known to persist only on a few seamounts. The main geomorphic features of the Chatham Rise are the smoothly sloping flanks (predominantly north- and south-facing), the flat-lying, though locally topographically irregular crestal region (including several prominent banks), and isolated groups of volcanic peaks or seamounts, with occasional sea valleys found on the flanks. Except for the emergent Chatham Islands, the rise deepens generally to the east, eventually merging with the abyssal plain of the Southwest Pacific Basin.

Seamount biodiversity: high variability both within and between seamounts in the Ross Sea region of Antarctica

Seamounts in the Ross Sea were surveyed during the New Zealand IPY-CAML research voyage in 2008. Admiralty seamount, and seamounts of the Scott Island Seamount chain (Scott Island, Scott A, Scott B, Scott South) were sampled to examine variability in benthic faunal communities of seamounts. Multivariate analysis of video and still image data showed that benthic community composition was highly variable both within and between seamounts. The distribution patterns of benthic communities varied considerably, with very high densities of certain taxa in some places, sparse fauna in many others, and highly patchy distributions for others. Dissimilarities between seamounts were influenced by differences in population densities of taxa, the proportions of transects in which taxa were present, and by taxa being restricted to only one seamount or group of seamounts. Scott Island seamount was less similar to the others, despite the greater distances between Admiralty and Scott A and B seamounts. The observed variability in community composition was mainly associated with differences in depth, chlorophyll concentration, and seabed rugosity. Results support an earlier hypothesis that the location of seamounts relative to the Ross Sea gyre could be an important factor determining the composition and abundance of benthic invertebrate communities.

Spatial patterns and environmental drivers of benthic infaunal community structure and ecosystem function on the New Zealand continental margin

To investigate regional drivers of spatial patterns in macro- and meio-faunal community structure (abundance, biomass and taxonomic diversity) and ecosystem function (sediment community oxygen consumption [SCOC]), we sampled two regions in close proximity on New Zealands continental margin—the Chatham Rise and the Challenger Plateau. Sites (n = 15) were selected in water depths ranging from 266–1212 m to generate a gradient in sedimentary properties and, in particular, surface pelagic productivity. Both macro- and meio-fauna abundance and biomass was 2–3.5 times higher on the Chatham Rise than on the Challenger Plateau, reflecting regional differences in pelagic primary production. We also found significant inter- and intra-regional differences in macro-fauna taxonomic diversity with two distinctive site groupings in each region. Univariate and multivariate measures of macro-fauna community attributes were most strongly correlated with sediment photosynthetic pigment (explaining 24%–59% of the variation). Sediment pigment content was as equally important in explaining meio-fauna community structure (36%–7%). Unlike community structure, SCOC was most strongly correlated with depth (44%), most likely reflecting temperature effects on benthic metabolism. Our results highlight the importance of a benthic labile food supply in structuring infaunal communities on continental margins and emphasise a tight coupling between pelagic and benthic habitats.

Habitat Differences in Deep-Sea Megafaunal Communities off New Zealand: Implications for Vulnerability to Anthropogenic Disturbance and Management

Research on benthic communities in the deep sea has focused largely on habitats in isolation, with few studies considering multiple habitats simultaneously in a comparable manner. The present study aimed to determine the structural differences in benthic communities of continental slope, seamount, canyon, vent, and seep habitats, and assess their relative vulnerabilities to disturbance from bottom trawling and potential seabed mining. Megafaunal invertebrate communities of these habitats were sampled in two regions off New Zealand, in four depth strata between 700 and 1500 m, using an epibenthic sled and a beam trawl. Patterns of community and trophic structure, and the potential influence of environmental variables, were determined using multivariate analyses. The difference in community structure between regions was greater than among habitats and depth strata. Levels of food availability may explain regional differences in community structure, although some influence of fishing disturbance is also possible. Differences in community and trophic structure were most pronounced between the chemosynthetic vent and seep habitats, and other habitats. Differences among these other habitats within a region were inconsistent, except that canyon and slope communities always differed from each other. Community and trophic structural patterns were partly explained by the environmental differences observed among habitats. The relative vulnerabilities of benthic communities to human disturbance in the two regions were determined based on patterns of abundance and feeding mode of the megafauna. Communities of vent and seep habitats were assessed to be more vulnerable to disturbance than those of the other habitats based on a number of habitat-related attributes. However, the relative vulnerability of megafaunal communities at slope, canyon, and seamount habitats could not confidently be assessed on a habitat basis alone. The results of the present study have implications for how regional and habitat differences in benthic communities are incorporated into spatial management options for the deep sea.

Evidence of niche conservatism and host fidelity in the polar shrimp Lebbeus kiae n. sp. (Decapoda: Caridea: Thoridae) from the Ross Sea, Antarctica

We report a remarkable case of ‘bipolarism’, where two different polar species, namely Lebbeus polaris in the northern hemisphere and Lebbeus kiae n. sp., here described from the Southern Ocean, have been found to share similar ecologies. Despite the great geographical distance between the two species, both show high host fidelity in associations with two congeneric sea anemones: Bolocera tuediae and Bolocera kerguelensis, respectively. A close molecular phylogenetic relationship between the two Lebbeus species is indicated by COI data, which clearly show them as sister clades with respect to other congeners as well as a plurality of other Antarctic species. This is the first reported case of a defensive association in the Southern Ocean involving shrimps and sea anemones. The distribution of the new species, limited to seamount systems off the Ross Sea, may be the result of a specific colonisation/speciation event in the past, although more molecular data are needed to unravel the phylogenetic relationships within the genus Lebbeus. Despite this uncertainty, the persistence of ecological traits, i.e., the defensive association with sea anemones, indicates the presence of niche conservatism in this clade of shrimps.