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The geochemistry of deep-sea coral skeletons: a review of vital effects and applications for palaeoceanography

Deep-sea corals were discovered over a century ago, but it is only over recent years that focused efforts have been made to explore the history of the oceans using the geochemistry of their skeletal remains. They offer a promising archive of past oceanic environments given their global distribution, layered growth patterns, longevity and preservation as well as our ability to date them using radiometric techniques. This paper provides an overview of the current state-of-the-art in terms of geochemical approaches to using deep-sea coral skeletons to explore the history of the ocean. Deep-sea coral skeletons have a wide array of morphologies (e.g. solitary cup corals, branching colonial corals) and materials (calcite, aragonite and proteins). As such their biomineralization strategies are diverse, leading to complex geochemistry within coral skeletons. Notwithstanding these complications, progress has been made on developing methods for reconstructing the oceanographic environment in the past using trace elements and isotopic methods. Promising approaches within certain coral groups include clumped isotopes and Mg/Li for temperature reconstructions, boron isotopes and radiocarbon for carbon cycling, εNd, and radiocarbon for circulation studies and δ 15 N, P/Ca and Ba/Ca for nutrient tracer studies. Likewise there is now a range of techniques for dating deep-sea corals skeletons (e.g. U-series, radiocarbon), and determining their growth rates (e.g. radiocarbon and 210 Pb). Dating studies on historic coral populations in the Atlantic, Southern Ocean and Pacific point to climate and environmental changes being dominant controls on coral populations over millennial and orbital timescales. This paper provides a review of a range of successes and promising approaches. It also highlights areas in which further research would likely provide new insights into biomineralization, palaeoceanography and distribution of past coral populations.

Deep-Sea Research Part II: Topical Studies in Ocea

Temporal and spatial distributions of cold-water corals in the Drake Passage: insights from the last 35,000 years

Scleractinian corals have a global distribution ranging from shallow tropical seas to the depths of the Southern Ocean. Although this distribution is indicative of the corals having a tolerance to a wide spectrum of environmental conditions, individual species seem to be restricted to a much narrower range of ecosystem variables. One way to ascertain the tolerances of corals, with particular focus on the potential impacts of changing climate, is to reconstruct their growth history across a range of environmental regimes. This study examines the spatial and temporal distribution of the solitary scleractinian corals Desmophyllum dianthus, Gardineria antarctica, Balanophyllia malouinensis, Caryophyllia spp. and Flabellum spp. from five sites in the Drake Passage which cross the major frontal zones. A rapid reconnaissance radiocarbon method was used to date more than 850 individual corals. Coupled with U-Th dating, an age range of present day back to more than 100 thousand years was established for corals in the region. Within this age range there are distinct changes in the temporal and spatial distributions of these corals, both with depth and latitude, and on millennial timescales. Two major patterns that emerge are: (1) D. dianthus populations show clear variability in their occurrence through time depending on the latitudinal position within the Drake Passage. North of the Subantarctic Front, D. dianthus first appears in the late deglaciation (~17,000 years ago) and persists to today. South of the Polar Front, in contrast, early deglacial periods, with a few modern occurrences. A seamount site between the two fronts exhibits characteristics similar to both the northern and southern sites. This shift across the frontal zones within one species cannot yet be fully explained, but it is likely to be linked to changes in surface productivity, subsurface oxygen concentrations, and carbonate saturation state. (2) at locations where multiple genera were dated, differences in age and depth distribution of the populations provide clear evidence that each genus has unique environmental requirements to sustain its population.

Deep-Sea Research Part II: Topical Studies in Ocea

Geomorphic characterization of four shelf-sourced submarine canyons along the U.S. Mid-Atlantic continental margin

Shelf-sourced submarine canyons are common features of continental margins and are fundamental to deep-sea sedimentary systems. Despite their geomorphic and geologic significance, relatively few passive margin shelf-breaching canyons worldwide have been mapped using modern geophysical methods. Between 2007 and 2012 a series of geophysical surveys was conducted across four major canyons of the US Mid-Atlantic margin: Wilmington, Baltimore, Washington, and Norfolk canyons. More than 5700 km 2 of high-resolution multibeam bathymetry and 890 line-km of sub-bottom CHIRP profiles were collected along the outer shelf and uppermost slope (depths of 80-1200 m). The data allowed us to compare and contrast the fine-scale morphology of each canyon system. The canyons have marked differences in the morphology and orientation of canyon heads, steepness and density of sidewall gullies, and the character of the continental shelf surrounding canyon rims. Down-canyon axial profiles for Washington, Baltimore and Wilmington canyons have linear shapes, and each canyon thalweg exhibits morphological evidence for recent, relatively small-scale sediment transport. For example, Washington Canyon displays extremely steep wall gradients and contains ~100 m wide, 5–10 m deep, v-shaped incisions down the canyon axis, suggesting modern or recent sediment transport. In contrast, the convex axial thalweg profile, the absence of thalweg incision, and evidence for sediment infilling at the canyon head, suggest that depositional processes strongly influence Norfolk Canyon during the current sea-level high-stand. The north walls of Wilmington, Washington and Norfolk canyons are steeper than the south walls due to differential erosion, though the underlying cause for this asymmetry is not clear. Furthermore, we speculate that most of the geomorphic features observed within the canyons (e.g., terraces, tributary canyons, gullies, and hanging valleys) were formed during the Pleistocene, and show only subtle modification by Holocene processes active during the present sea-level high-stand.

Cape Hatteras;Georges Bank;Hudson Canyon

Alexandrium fundyense cysts in the Gulf of Maine: long-term time series of abundance and distribution, and linkages to past and future blooms

Here we document Alexandrium fundyense cyst abundance and distribution patterns over nine years (1997 and 2004–2011) in the coastal waters of the Gulf of Maine (GOM) and identify linkages between those patterns and several metrics of the severity or magnitude of blooms occurring before and after each autumn cyst survey. We also explore the relative utility of two measures of cyst abundance and demonstrate that GOM cyst counts can be normalized to sediment volume, revealing meaningful patterns equivalent to those determined with dry weight normalization. Cyst concentrations were highly variable spatially. Two distinct seedbeds (defined here as accumulation zones with>300 cysts cm −3 ) are evident, one in the Bay of Fundy (BOF) and one in mid-coast Maine. Overall, seedbed locations remained relatively constant through time, but their area varied 3–4 fold, and total cyst abundance more than 10 fold among years. A major expansion of the mid-coast Maine seedbed occurred in 2009 following an unusually intense A. fundyense bloom with visible red-water conditions, but that feature disappeared by late 2010. The regional system thus has only two seedbeds with the bathymetry, sediment characteristics, currents, biology, and environmental conditions necessary to persist for decades or longer. Strong positive correlations were confirmed between the abundance of cysts in both the 0–1 and the 0–3 cm layers of sediments in autumn and geographic measures of the extent of the bloom that occurred the next year (i.e., cysts→blooms), such as the length of coastline closed due to shellfish toxicity or the southernmost latitude of shellfish closures. In general, these metrics of bloom geographic extent did not correlate with the number of cysts in sediments following the blooms (blooms→cysts). There are, however, significant positive correlations between 0–3 cm cyst abundances and metrics of the preceding bloom that are indicative of bloom intensity or vegetative cell abundance (e.g., cumulative shellfish toxicity, duration of detectable toxicity in shellfish, and bloom termination date). These data suggest that it may be possible to use cyst abundance to empirically forecast the geographic extent of the forthcoming bloom and, conversely, to use other metrics from bloom and toxicity events to forecast the size of the subsequent cyst population as the inoculum for the next year's bloom. This is an important step towards understanding the excystment/encystment cycle in A. fundyense bloom dynamics while also augmenting our predictive capability for this HAB-forming species in the GOM.

Maine

Investigating the importance of sediment resuspension in Alexandrium fundyense cyst population dynamics in the Gulf of Maine

Cysts of Alexandrium fundyense , a dinoflagellate that causes toxic algal blooms in the Gulf of Maine, spend the winter as dormant cells in the upper layer of bottom sediment or the bottom nepheloid layer and germinate in spring to initiate new blooms. Erosion measurements were made on sediment cores collected at seven stations in the Gulf of Maine in the autumn of 2011 to explore if resuspension (by waves and currents) could change the distribution of over-wintering cysts from patterns observed in the previous autumn; or if resuspension could contribute cysts to the water column during spring when cysts are viable. The mass of sediment eroded from the core surface at 0.4 Pa ranged from 0.05 kg m −2 near Grand Manan Island, to 0.35 kg m −2 in northern Wilkinson Basin. The depth of sediment eroded ranged from about 0.05 mm at a station with sandy sediment at 70 m water depth on the western Maine shelf, to about 1.2 mm in clayey–silt sediment at 250 m water depth in northern Wilkinson Basin. The sediment erodibility measurements were used in a sediment-transport model forced with modeled waves and currents for the period October 1, 2010 to May 31, 2011 to predict resuspension and bed erosion. The simulated spatial distribution and variation of bottom shear stress was controlled by the strength of the semi-diurnal tidal currents, which decrease from east to west along the Maine coast, and oscillatory wave-induced currents, which are strongest in shallow water. Simulations showed occasional sediment resuspension along the central and western Maine coast associated with storms, steady resuspension on the eastern Maine shelf and in the Bay of Fundy associated with tidal currents, no resuspension in northern Wilkinson Basin, and very small resuspension in western Jordan Basin. The sediment response in the model depended primarily on the profile of sediment erodibility, strength and time history of bottom stress, consolidation time scale, and the current in the water column. Based on analysis of wave data from offshore buoys from 1996 to 2012, the number of wave events inducing a bottom shear stress large enough to resuspend sediment at 80 m ranged from 0 to 2 in spring (April and May) and 0 to 10 in winter (October through March). Wave-induced resuspension is unlikely in water greater than about 100 m deep. The observations and model results suggest that a millimeter or so of sediment and associated cysts may be mobilized in both winter and spring, and that the frequency of resuspension will vary interannually. Depending on cyst concentration in the sediment and the vertical distribution in the water column, these events could result in a concentration in the water column of at least 10 4 cysts m −3 . In some years, resuspension events could episodically introduce cysts into the water column in spring, where germination is likely to be facilitated at the time of bloom formation. An assessment of the quantitative effects of cyst resuspension on bloom dynamics in any particular year requires more detailed investigation.

Bay Of Fundy;Grand Manan Island;Gulf Of Maine;Jord

Near-bottom circulation and dispersion of sediment containing Alexandrium fundyense cysts in the Gulf of Maine during 2010-2011

The life cycle of Alexandrium fundyense in the Gulf of Maine includes a dormant cyst stage that spends the winter predominantly in the bottom sediment. Wave-current bottom stress caused by storms and tides induces resuspension of cyst-containing sediment during winter and spring. Resuspended sediment could be transported by water flow to different locations in the Gulf and the redistribution of sediment containing A. fundyense cysts could alter the spatial and temporal manifestation of its spring bloom. The present study evaluates model near-bottom flow during storms, when sediment resuspension and redistribution are most likely to occur, between October and May when A. fundyense cells are predominantly in cyst form. Simulated water column sediment (mud) concentrations from representative locations of the Gulf are used to initialize particle tracking simulations for the period October 2010–May 2011. Particles are tracked in full three-dimensional model solutions including a sinking velocity characteristic of cyst and aggregated mud settling (0.1 mm s − 1). Although most of the material was redeposited near the source areas, small percentages of total resuspended sediment from some locations in the western (~4%) and eastern (2%) Maine shelf and the Bay of Fundy (1%) traveled distances longer than 100 km before resettling. The redistribution changed seasonally and was sensitive to the prescribed sinking rate. Estimates of the amount of cysts redistributed with the sediment were small compared to the inventory of cysts in the upper few centimeters of sediment but could potentially have more relevance immediately after deposition.

Maine

Growth rates and ages of deep-sea corals impacted by the Deepwater Horizon oil spill

The impact of the April 2010 Deepwater Horizon (DWH) spill on deep-sea coral communities in the Gulf of Mexico (GoM) is still under investigation, as is the potential for these communities to recover. Impacts from the spill include observation of corals covered with flocculent material, with bare skeleton, excessive mucous production, sloughing tissue, and subsequent colonization of damaged areas by hydrozoans. Information on growth rates and life spans of deep-sea corals is important for understanding the vulnerability of these ecosystems to both natural and anthropogenic perturbations, as well as the likely duration of any observed adverse impacts. We report radiocarbon ages and radial and linear growth rates based on octocorals ( Paramuricea spp. and Chrysogorgia sp.) collected in 2010 and 2011 from areas of the DWH impact. The oldest coral radiocarbon ages were measured on specimens collected 11 km to the SW of the oil spill from the Mississippi Canyon (MC) 344 site: 599 and 55 cal yr BP, suggesting continuous life spans of over 600 years for Paramuricea biscaya, the dominant coral species in the region. Calculated radial growth rates, between 0.34 μm yr −1 and 14.20 μm yr −1 , are consistent with previously reported proteinaceous corals from the GoM. Anomalously low radiocarbon (Δ 14 C) values for soft tissue from some corals indicate that these corals were feeding on particulate organic carbon derived from an admixture of modern surface carbon and a low 14 C carbon source. Results from this work indicate fossil carbon could contribute 5–10% to the coral soft tissue Δ 14 C signal within the area of the spill impact. The influence of a low 14C carbon source (e.g., petro-carbon) on the particulate organic carbon pool was observed at all sites within 30 km of the spill site, with the exception of MC118, which may have been outside of the dominant northeast-southwest zone of impact. The quantitatively assessed extreme longevity and slow growth rates documented here highlight the vulnerability of these long-lived deep sea coral species to disturbance.

Deep-Sea Research Part II: Topical Studies in Ocea

Low incidence of clonality in cold water corals revealed through the novel use of standardized protocol adapted to deep sea sampling

Sampling in the deep sea is a technical challenge, which has hindered the acquisition of robust datasets that are necessary to determine the fine-grained biological patterns and processes that may shape genetic diversity. Estimates of the extent of clonality in deep-sea species, despite the importance of clonality in shaping the local dynamics and evolutionary trajectories, have been largely obscured by such limitations. Cold-water coral reefs along European margins are formed mainly by two reef-building species, Lophelia pertusa and Madrepora oculata . Here we present a fine-grained analysis of the genotypic and genetic composition of reefs occurring in the Bay of Biscay, based on an innovative deep-sea sampling protocol. This strategy was designed to be standardized, random, and allowed the georeferencing of all sampled colonies. Clonal lineages discriminated through their Multi-Locus Genotypes (MLG) at 6–7 microsatellite markers could thus be mapped to assess the level of clonality and the spatial spread of clonal lineages. High values of clonal richness were observed for both species across all sites suggesting a limited occurrence of clonality, which likely originated through fragmentation. Additionally, spatial autocorrelation analysis underlined the possible occurrence of fine-grained genetic structure in several populations of both L. pertusa and M. oculata . The two cold-water coral species examined had contrasting patterns of connectivity among canyons, with among-canyon genetic structuring detected in M. oculata , whereas L. pertusa was panmictic at the canyon scale. This study exemplifies that a standardized, random and georeferenced sampling strategy, while challenging, can be applied in the deep sea, and associated benefits outlined here include improved estimates of fine grained patterns of clonality and dispersal that are comparable across sites and among species.

Deep-Sea Research Part II: Topical Studies in Ocea

Macrofaunal communities associated with chemosynthetic habitats from the U.S. Atlantic margin: A comparison among depth and habitat types

Hydrocarbon seeps support distinct benthic communities capable of tolerating extreme environmental conditions and utilizing reduced chemical compounds for nutrition. In recent years, several locations of methane seepage have been mapped along the U.S. Atlantic continental slope. In 2012 and 2013, two newly discovered seeps were investigated in this region: a shallow site near Baltimore Canyon (BCS, 366–412 m) and a deep site near Norfolk Canyon (NCS, 1467–1602 m), with both sites containing extensive chemosynthetic mussel bed and microbial mat habitats. Sediment push cores, suction samples, and Ekman box cores were collected to quantify the abundance, diversity, and community structure of benthic macrofauna (>300 μm) in mussel beds, mats, and slope habitats at both sites. Community data from the deep site were also assessed in relation to the associated sediment environment (organic carbon and nitrogen, stable carbon and nitrogen isotopes, grain size, and depth). Infaunal assemblages and densities differed both between depths and among habitat types. Macrofaunal densities in microbial mats were four times greater than those present in mussel beds and slope sediments and were dominated by the annelid families Dorvilleidae, Capitellidae, and Tubificidae, while mussel habitats had higher proportions of crustaceans. Diversity was lower in BCS microbial mat habitats, but higher in mussel and slope sediments compared to NCS habitats. Multivariate statistical analysis revealed specific sediment properties as important for distinguishing the macrofaunal communities, including larger grain sizes present within NCS microbial mat habitats and depleted stable carbon isotopes (δ 13 C) in sediments present at mussel beds. These results suggest that habitat differences in the quality and source of organic matter are driving the observed patterns in the infaunal assemblages, including high β diversity and high variability in the macrofaunal community composition. This study is the first investigation of seep infauna along the U.S. Atlantic slope north of the Blake Ridge Diapir and provides a baseline for future regional comparisons to other seep habitats along the Atlantic margin.

Deep-Sea Research Part II: Topical Studies in Ocea

A phylogenetic perspective on diversity of Galatheoidea ( Munida, Munidopsis ) from cold-water coral and cold seep communities in the western North Atlantic Ocean

Squat lobsters (Galatheoidea and Chirostyloidea), a diverse group of decapod crustaceans, are ubiquitous members of the deep-sea fauna. Within Galatheoidea, the genera Munida and Munidopsis are the most diverse, but accurate estimates of biodiversity are difficult due to morphological complexity and cryptic diversity. Four species of Munida and nine species of Munidopsis from cold-water coral (CWC) and cold seep communities in the northwestern Atlantic Ocean (NWA) and the Gulf of Mexico (GOM) were collected over eleven years and fifteen research cruises in order to assess faunal associations and estimate squat lobster biodiversity. Identification of the majority of specimens was determined morphologically. Mitochondrial COI sequence data, obtained from material collected during these research cruises, was supplemented with published sequences of congeners from other regions. The phylogenetic analysis of Munida supports three of the four NWA and GOM species ( M. microphthalma , M. sanctipauli , and M. valida ) as closely related taxa. The fourth species, Munida iris , is basal to most other species of Munida , and is closely related to M. rutllanti, a species found in the northeastern Atlantic Ocean (NEA). The majority of the nine species of Munidopsis included in our analyses were collected from chemosynthetic cold seep sites from the GOM. While seep taxa were scattered throughout the phylogenetic tree, four of these species ( Munidopsis livida , M. similis, M. bermudezi , and M. species A ) from the NWA and the GOM were part of a large eighteen-species clade that included species collected from Pacific Ocean chemosynthetic habitats, such as hydrothermal vents and whale falls. Shinkaia crosnieri was the sister taxon to the chemosynthetic clade, and M. livida was the most basal member of this clade. Munidopsis sp. B , an undescribed species with representative individuals collected from two GOM chemosynthetic sites, exhibited the largest genetic distance from other northern Atlantic species. Generally, intraspecific diversity was lower and patterns of haplotype diversity more simple in species of Munidopsis relative to Munida . This study puts two genera of NWA and GOM squat lobsters into a population genetic and phylogenetic context with regard to biogeography and habitat to enhance understanding of the history and evolutionary trajectories of these morphologically and ecologically diverse groups.

Deep-Sea Research Part II: Topical Studies in Ocea

Gene transcription patterns in response to low level petroleum contaminants in Mytilus trossulus from field sites and harbors in southcentral Alaska

The 1989 Exxon Valdez oil spill damaged a wide range of natural resources, including intertidal communities, and post-spill studies demonstrated acute and chronic exposure and injury to an array of species. Standard toxicological methods to evaluate petroleum contaminants have assessed tissue burdens, with fewer assays providing indicators of health or physiology, particularly when contaminant levels are low and chronic. Marine mussels are a ubiquitous and crucial component of the nearshore environment, and new genomic technologies exist to quantify molecular responses of individual mussels to stimuli, including exposure to polycyclic aromatic hydrocarbons (PAHs). We used gene-based assays of exposure and physiological function to assess chronic oil contamination using the Pacific blue mussel, Mytilus trossulus . We developed a diagnostic gene transcription panel to investigate exposure to PAHs and other contaminants and its effects on mussel physiology and health. During 2012–2015, we analyzed mussels from five field sites in western Prince William Sound, Alaska, with varying oil histories from the 1989 Exxon Valdez oil spill, and from three boat harbors in the area. Gene transcription patterns of mussels from harbors were consistent with elevated exposure to PAHs or other contaminants, whereas transcription patterns of mussels sampled from shorelines in areas affected by the oil spill indicated no PAH exposure.

Alaska

Seafloor fluid seeps on Kimki Ridge, offshore southern California: Links to active strike-slip faulting

The Kimki Ridge fluid seeps are located in western Catalina Basin about 60 km southwest of the southern California mainland and at a water depth of approximately 1100 m. Multichannel seismic reflection profiles collected by the U.S. Geological Survey (USGS) in 2014 show acoustic transparency within the Kimki Ridge, suggesting the possibility of fluid seeps and possible sub-seafloor fluid pathways. Subsequent multibeam bathymetric and backscatter intensity data collected during a cooperative University of Washington/USGS cruise in early 2016 show subtle seafloor buildups with high acoustic backscatter (reflectivity) in three places along Kimki Ridge, supporting the existence of fluid seepage. A Remotely Operated Vehicle (ROV) dive, conducted as part of the Nautilus Exploration Program, took place in August 2016 to confirm the presence of these previously unknown seeps and document their characteristics as well as those of any associated biological communities. Two of the three seeps were explored by ROV, and showed abundant evidence of fluid seepage, including characteristic algal mats, chemosynthetic clams, and authigenic carbonate formation. The seeps are comprised of carbonate buildups 1–3 m thick and 300–500 m across. Within these areas, we interpret broad crater-like depressions 30–50 m across and 1–2 m deep to be individual seep vents. The seep areas appear to be broad zones of diffuse seepage that support chemosynthetic biologic communities; however, active venting was not observed. Geochemical analyses of rock samples collected from the seeps indicate microbially driven anaerobic oxidation of methane at or near the sediment water interface. Seismic-reflection profiles show chimney-like fluid pathways along the limbs and in the axis of the fold forming Kimki Ridge, and evidence of methane in shallow sediments can be traced into the adjacent Catalina Basin. A system of closely spaced faults located at the axis of the Kimki Ridge anticline may serve as pathways to allow fluid flow to the seafloor. Our data are consistent with other studies that suggest that transpression is an important component in the formation and localization of fluid seeps in a strike-slip setting, implying that seep formation may be a common occurrence at fault stepovers or transpressional bends in strike-slip systems.

California

Ancient impact structures on modern continental shelves: The Chesapeake Bay, Montagnais, and Toms Canyon craters, Atlantic margin of North America

Three ancient impact craters (Chesapeake Bay - 35.7 Ma; Toms Canyon - 35.7 Ma; Montagnais - 51 Ma) and one multiring impact basin (Chicxulub - 65 Ma) are currently known to be buried beneath modern continental shelves. All occur on the passive Atlantic margin of North America in regions extensively explored by seismic reflection surveys in the search for oil and gas reserves. We limit our discussion herein to the three youngest structures. These craters were created by submarine impacts, which produced many structural and morphological features similar in construction, composition, and variability to those documented in well-preserved subaerial and planetary impact craters. The subcircular Chesapeake Bay (diameter 85 km) and ovate Montagnais (diameter 45-50 km) structures display outer-rim scarps, annular troughs, peak rings, inner basins, and central peaks similar to those incorporated in the widely cited conceptual model of complex impact craters. These craters differ in several respects from the model, however. For example, the Montagnais crater lacks a raised lip on the outer rim, the Chesapeake Bay crater displays only small remnants of a raised lip, and both craters contain an unusually thick body of impact breccia. The subtriangular Toms Canyon crater (diameter 20-22 km), on the other hand, contains none of the internal features of a complex crater, nor is it typical of a simple crater. It displays a prominent raised lip on the outer rim, but the lip is present only on the western side of the crater. In addition, each of these craters contains some distinct features, which are not present in one or both of the others. For example, the central peak at Montagnais rises well above the elevation of the outer rim, whereas at Chesapeake Bay, the outer rim is higher than the central peak. The floor of the Toms Canyon crater is marked by parallel deep troughs and linear ridges formed of sedimentary rocks, whereas at Chesapeake Bay, the crater floor contains concentric faults and compression ridges formed in rocks of the crystalline basement. The Chesapeake Bay crater is distinguished further by its cluster of at least 23 adjacent secondary craters. The North American tektite strewn field, a widespread deposit of distal ejecta, is thought to be derived from the Chesapeake Bay impact, perhaps with a small contribution from the Toms Canyon impact. No ejecta field is known to be associated with the Montagnais impact. No immediate major extinction event is directly linked to any of these three impacts. There is evidence, however, that the Chesapeake Bay and Toms Canyon impacts helped initiate a long-term pulse of warm global climate, whose eventual dissipation coincided with an early Oligocene mass extinction event, 2 Ma after the impacts.

Deep-Sea Research Part II: Topical Studies in Ocea

Natural radionuclides and plutonium in sediments from the western Arctic Ocean: Sedimentation rates and pathways of radionuclides

Sediment cores collected during R.V. Polar Sea AOS94 expedition from the Chukchi Shelf to the North Poke were analyzed for several decay-series natural radionuclides and Pu isotopes to study sedimentation rates and pathways of radionuclides in the western Arctic Ocean. The measured sedimentation rates vary by more than three orders of magnitude along the transect, from 210Pb based rates of 200-700 cm kyr-1 over the Chukchi Shelf and 89 cm kyr-1 at the Chukchi Slope to 230Th-based rates of 0.02-0.3 cm kyr-1 at various settings in the deep basin. 230Th(ex) profiles in the central western Arctic Basin are characterized by a cyclic pattern and a pronounced sub-surface maximum superimposed on an overall decrease with depth. Sediment inventories of excess 210Pb and 230Th in the deep basin as a whole cannot account for their in situ production and 2610Pb fall-out. The opposite is true at the slope and shallower waters. We contend that, as with other ocean basins, boundary scavenging also exists in the Arctic Ocean. The broad continental shelves and the slope region may have the potential of removing all or moat of the particle-reactive radionuclides unaccounted for in the deep basin. The Pu isotope data are consistent with the notion of boundary scavenging. Sediment inventories and concentrations of Pu decrease rapidly offshore. Isotopic composition of Pu suggests mixing of fall-out Pu, which decreases with increasing latitudes, and fuels reprocessing Pu derived from the Russian and Atlantic sides of the Arctic Ocean. Although fuel reprocessing Pu has impinged on the Chukchi Slope, its existence over the Chukchi Shelf is not evident and probably overshadowed by fall-out Pu.

Deep-Sea Research Part II: Topical Studies in Ocea

Enumeration of viruses and prokaryotes in deep-sea sediments and cold seeps of the Gulf of Mexico

Little is known about the distribution and abundance of viruses in deep-sea cold-seep environments. Like hydrothermal vents, seeps support communities of macrofauna that are sustained by chemosynthetic bacteria. Sediments close to these communities are hypothesized to be more microbiologically active and therefore to host higher numbers of viruses than non-seep areas. Push cores were taken at five types of Gulf of Mexico habitats at water depths below 1000 m using a remotely operated vehicle (ROV). The habitats included non-seep reference sediment, brine seeps, a microbial mat, an urchin field, and a pogonophoran worm community. Samples were processed immediately for enumeration of viruses and prokaryotes without the addition of a preservative. Prokaryote counts were an order of magnitude lower in sediments directly in contact with macrofauna (urchins, pogonophorans) compared to all other samples (10 7 vs. 10 8 cells g -1 dry weight) and were highest in areas of elevated salinity (brine seeps). Viral-Like Particle (VLP) counts were lowest in the reference sediments and pogonophoran cores (10 8 VLP g -1 dry wt), higher in brine seeps (10 9 VLP g -1 dry wt), and highest in the microbial mats (10 10 VLP g -1 dry wt). Virus-prokaryote ratios (VPR) ranged from <5 in the reference sediment to >30 in the microbial mats and >60 in the urchin field. VLP counts and VPR were all significantly greater than those reported from sediments in the deep Mediterranean Sea and in most cases were higher than recent data from a cold-seep site near Japan. The high VPR suggest that greater microbial activity in or near cold-seep environments results in greater viral production and therefore higher numbers of viruses.

Gulf Of Mexico;Green Canyon;Atwater Valley;Alamino

Characterizing the sponge grounds of Grays Canyon, Washington, USA

Deep-sea sponge grounds are relatively understudied ecosystems that may provide key habitats for a large number of fish and invertebrates including commercial species. Glass sponge grounds have been discovered from the tropics to polar regions but there are only a few places with high densities of dictyonine sponges. Dictyonine glass sponges have a fused skeleton, which stays intact when they die and in some areas the accumulation of successive generations of sponges leads to the formation of reefs. In 2010 and 2016, we surveyed an area near Grays Canyon in Washington, USA, where dense aggregations of glass sponges and potential sponge reefs were discovered in 2007. Our primary aims were to make a preliminary assessment of whether the glass sponges form reefs at this location, characterize the sponge assemblage present at this site and examine associations between the sponges and commercially important species. Multibeam mapping and sub-bottom profiling indicate that the glass sponges at this site do not form reefs and are mostly attached to hard substrates. Analysis of photographs collected by an autonomous underwater vehicle and samples collected by a remotely operated vehicle guided by telepresence revealed the presence of two abundant dictyonine sponge species at this site, Heterochone calyx and Aphrocallistes vastus (mean densities = 1.43 ± 0.057 per 10 m2, max = 24 per 10 m2). We also observed a large number of non-reef-building glass sponges and various demosponges including a potentially new species in the genus Acarnus. A diverse fish assemblage was recorded at this site including eight species of rockfish. Rockfish abundance was positively related to sponge abundance. Spot prawns (Pandalus platyceros) were also abundant and were strongly associated with sponges. Despite not finding sponge reefs, this is an ecologically significant area. Further research is necessary to determine the environmental factors that give rise to the abundance of large dictyonine sponges at this location and also to determine if other similar sponge grounds exist along the west coast of the United States.

Washington

A comparison of deep-sea coral and sponge species distribution models and the impact of ocean currents in the Southern California Bight

Deep-sea corals and sponges (DSCS) are signature taxa of deep-water habitats, but ecological mechanisms that drive their geographic distributions can be difficult to uncover due to the challenges of surveying deep-water ecosystems and limited oceanographic data. A comparison of species distribution models was made for three DSCS ( Antipathes dendochristos, Plumarella longispina, and an unidentified sponge morphotype known as Porifera #2), two of which are associated with young-of-the-year and adult rockfish taxa ( Sebastes spp.). Models were built using generalized additive models (GAMs) that accounted for spatial autocorrelation. These models were compared to Maxent models, a commonly used method. All models utilized bathymetrically derived variables (including depth, slope, and Bathymetric Position Index) and output from a coupled physical-biogeochemical ocean model (including current direction and magnitude, food abundance, and dissolved oxygen). Direction of benthic ocean currents played an important role in predictions, pointing to large-scale ecological processes that may determine presence of DSCS. GAMs outperformed Maxent models and predicted more ecologically interpretable patterns. Additionally we quantified the predicted amount of suitable habitat that is currently located within some protected areas of the Southern California Bight and show that many hot spots occur outside protected areas. Given the importance of DSCS for multiple demersal fish species of commercial or conservation concerns, this research provides valuable information that natural resource managers can use as a tool in the Southern California Bight.

California

Megafaunal community structure on ferromanganese and phosphorite hardgrounds in the Southern California Borderland

The Southern California Borderland (SCB) is a topographically complex region on the active continental margin that hosts varied hardground habitats, including ferromanganese (FeMn) crusts and phosphorites, marine minerals being considered for resource extraction. The SCB is influenced by seasonal upwelling and terrestrial inputs, and has a well-defined oxygen-minimum zone (OMZ). We analyzed megafaunal community composition, density, and diversity across 41 ROV video transects at 10 SCB hard substrate sites spanning depth (378–2765 m), temperature (1.79–7.96 °C), and oxygen (3.91–105.67 μM) gradients, with varying rock types (FeMn crusts, phosphorites, other rocks). We counted 32,426 individuals representing 146 unique taxa over a total area of 21,935 m 2 . Echinodermata contributed 48 % of the total abundance; Cnidaria 24 %; Porifera 13 %; Annelida 6 %; Arthropoda 6 %; Chordata 2 %; Mollusca, Foraminifera, and Hemichordata <1 %. Megafauna communities showed clear heterogeneity, with density, diversity, and community composition varying among transects within sites and across sites. Rock type recorded no influence on megafaunal density, but habitats that included FeMn crusts had higher diversity and distinct taxonomic composition. Location, depth, and oxygen were the most important variables explaining variance among megafaunal communities, with distinct communities formed at deeper depths and within the OMZ. This study provides baseline information about the megafauna inhabiting SCB hardgrounds, particularly those associated with minerals considered for their resource potential. It extends existing documentation of such communities to greater depths, reveals increased representation of sponges and corals below 1000 m, and highlights the variability within and among deep-sea megafaunal communities, offering context for informed policy decisions.

Southern California borderland