Xyloplax is a genus of three species of sea stars previously found only on sunken wood in the deep ocean. Their circular and petaloid bodies, which lend them their common name “sea daisy”, and their presumed exclusive diet of wood make them an unusual and rare element of deep-sea ecosystems. We describe here the fourth species of Xyloplax from the eastern Pacific Ocean, Xyloplax princealberti n. sp., which ranges from offshore Canada to the Gulf of California (Mexico) and Costa Rica. Though sampled geographically close to another described species of Xyloplax from the northeastern Pacific, X. janetae, this new species is unique morphologically and according to available DNA data. The short abactinal spines are the most obvious feature that distinguishes X. princealberti n. sp. from other Xyloplax. The minimum distance for mitochondrial cytochrome c oxidase subunit I from Xyloplax princealberti n. sp. to the only other available Xyloplax, X. janetae, was 13.5%. We also describe Ridgeia vestimentiferan tubeworm bushes from active hydrothermal vents as a new Xyloplax habitat, the first record of a non-wood substrate, and a new reproductive strategy, simultaneous hermaphroditism, for this genus. We generated the first mitochondrial genome for a member of Xyloplax and analyzed it with other available asteroid data using nucleotide-coding or amino acid (for protein-coding genes) plus nucleotide coding (for rRNA genes). The nucleotide-coding results place Xylopax as part of the clade Velatida, consistent with a previous phylogenomic analysis that included Xyloplax princealberti n. sp. (as Xyloplax sp.), though the placement of Velatida within Asteroidea differed. The amino acid plus nucleotide coding recovered Velatida to be a grade with X. princealberti n. sp. as sister group to all other Asteroidea.

Species inhabiting deep-sea hydrothermal vents are strongly influenced by the geological setting, as it provides the chemical-rich fluids supporting the food web, creates the patchwork of seafloor habitat, and generates catastrophic disturbances that can eradicate entire communities. The patches of vent habitat host a network of communities (a metacommunity) connected by dispersal of planktonic larvae. The dynamics of the metacommunity are influenced not only by birth rates, death rates and interactions of populations at the local site, but also by regional influences on dispersal from different sites. The connections to other communities provide a mechanism for dynamics at a local site to affect features of the regional biota. In this paper, we explore the challenges and potential benefits of applying metacommunity theory to vent communities, with a particular focus on effects of disturbance. We synthesize field observations to inform models and identify data gaps that need to be addressed to answer key questions including: (1) what is the influence of the magnitude and rate of disturbance on ecological attributes, such as time to extinction or resilience in a metacommunity; (2) what interactions between local and regional processes control species diversity, and (3) which communities are “hot spots” of key ecological significance. We conclude by assessing our ability to evaluate resilience of vent metacommunities to human disturbance (e.g., deep-sea mining). Although the resilience of a few highly disturbed vent systems in the eastern Pacific has been quantified, these values cannot be generalized to remote locales in the western Pacific or mid Atlantic where disturbance rates are different and information on local controls is missing.

Size, growth, and density have been studied for North American Pacific coast sea urchins Strongylocentrotus purpuratus, S. droebachiensis, S. polyacanthus, Mesocentrotus (Strongylocentrotus) franciscanus, Lytechinus pictus, Centrostephanus coronatus, and Arbacia stellata by various workers at diverse sites and for varying lengths of time from 1956 to present. Numerous peer-reviewed publications have used some of these data but some data have appeared only in graduate theses or the gray literature. There also are data that have never appeared outside original data sheets. Motivation for studies has included fisheries management and environmental monitoring of sewer and power plant outfalls as well as changes associated with disease epidemics. Studies also have focused on kelp restoration, community effects of sea otters, basic sea urchin biology, and monitoring. The data sets presented here are a historical record of size, density, and growth for a common group of marine invertebrates in intertidal and nearshore environments that can be used to test hypotheses concerning future changes associated with fisheries practices, shifts of predator distributions, climate and ecosystem changes, and ocean acidification along the Pacific Coast of North America and islands of the north Pacific. No copyright restrictions apply. Please credit this paper when using the data.

The deep sea encompasses the largest ecosystems on Earth. Although poorly known, deep seafloor ecosystems provide services that are vitally important to the entire ocean and biosphere. Rising atmospheric greenhouse gases are bringing about significant changes in the environmental properties of the ocean realm in terms of water column oxygenation, temperature, pH and food supply, with concomitant impacts on deep-sea ecosystems. Projections suggest that abyssal (3000–6000 m) ocean temperatures could increase by 1°C over the next 84 years, while abyssal seafloor habitats under areas of deep-water formation may experience reductions in water column oxygen concentrations by as much as 0.03 mL L–1 by 2100. Bathyal depths (200–3000 m) worldwide will undergo the most significant reductions in pH in all oceans by the year 2100 (0.29 to 0.37 pH units). O2 concentrations will also decline in the bathyal NE Pacific and Southern Oceans, with losses up to 3.7% or more, especially at intermediate depths. Another important environmental parameter, the flux of particulate organic matter to the seafloor, is likely to decline significantly in most oceans, most notably in the abyssal and bathyal Indian Ocean where it is predicted to decrease by 40–55% by the end of the century. Unfortunately, how these major changes will affect deep-seafloor ecosystems is, in some cases, very poorly understood. In this paper, we provide a detailed overview of the impacts of these changing environmental parameters on deep-seafloor ecosystems that will most likely be seen by 2100 in continental margin, abyssal and polar settings. We also consider how these changes may combine with other anthropogenic stressors (e.g., fishing, mineral mining, oil and gas extraction) to further impact deep-seafloor ecosystems and discuss the possible societal implications.

Authigenic carbonate rocks at methane seeps are recognized as hosting diverse and abundant invertebrate assemblages, with potential forcing from fluid seepage and hydrography. Mensurative studies of carbonate macrofauna (>0.3mm) at Hydrate Ridge, OR revealed little effect of water depth and overlying oxygenation (at 600m and 800m) but a large influence of seepage activity on density, taxonomic composition, diversity, and biological traits (feeding, lifestyle, motility, size and calcification). Rocks exposed to active seepage had 3–4× higher total macrofaunal densities than under inactive conditions. Assemblages exhibited higher species richness and reduced evenness (greater dominance) under active seepage than inactive conditions, but no difference in H′ or rarefaction diversity. Actively seeping sites were characterized by errant (motile), bacterial grazing, small- and medium-sized, heavily calcified species, whereas inactive sites exhibited a greater diversity of feeding modes and more burrowers, sessile, large and lightly calcified species. Active rocks supported more exogonid (Syllidae), ampharetid, and cirratulid polychaetes, provannid snails, pyropeltid limpets, nemerteans, and sponges; whereas inactive rocks supported higher densities of ophiuroids, isopods, gammarid amphipods, hydroids, Typosyllis (Syllidae) and tanaids. Transplant experiments, in which rocks were transferred between active and inactive sites at Hydrate Ridge North (600m), revealed that assemblages respond within 13months to increase or cessation of seepage, taking on the feeding, size and calcification characteristics of the background fauna at the new site. Lifestyles and motility patterns shifted more slowly as the sessile, attached species did not track seepage as quickly. Provannid snails and pyropeltid limpets rapidly colonized rocks transplanted to active sites and disappeared when transplanted to inactive sites. Given the known variability of fluid fluxes and rapid community response, a mosaic of communities changing in space and time is hypothesized to generate the relatively high species diversity at methane seeps.

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.

By creating novel habitats, habitat-modifying species can alter patterns of diversity and abundance in marine communities. Many sea urchins are important habitat modifiers in tropical and temperate systems. By eroding rocky substrata, urchins can create a mosaic of urchin-sized cavities or pits separated by exposed, often flat surfaces. These microhabitats appear to harbor distinct assemblages of species. We investigated how a temperate rocky intertidal community uses three small-scale (<100cm(2)) microhabitats created by or adjacent to populations of the purple sea urchin (Strongylocentrotus purpuratus): pits occupied by urchins, unoccupied pits, and adjacent flat spaces. In tidepools, flat spaces harbored the highest percent cover of algae and sessile fauna, followed by empty pits and then occupied pits. The Shannon diversity and richness of these sessile taxa were significantly higher in flat spaces and empty pits than in occupied pits. The composition of these primary space holders in the microhabitats also varied. Unlike primary space holders, mobile fauna exhibited higher diversity and richness in empty pits than in flat spaces and occupied pits, although results were not significant. The protective empty pit microhabitat harbored the highest densities of most trophic functional groups. Herbivores, however, were densest in flat spaces, concordant with high algal coverage. These results suggest the habitats created by S.purpuratus in addition to its biological activities alter community structure at spatial scales finer than those typically considered for sea urchins.

The recent discovery of a methane seep with diverse microhabitats and abundant groundfish in the San Diego Trough (1020 m) off the coast of Del Mar, California raised questions about the role of seep ecosystem functions and services in relation to continental margins. We used multicorer and ROV grab samples and an ROV survey to characterize macrofaunal structure, diversity, and trophic patterns in soft sediments and authigenic carbonates; seep microhabitats and taxa observed; and the abundance and spatial patterns of fishery-relevant species. Biogenic microhabitats near the Del Mar Seep included microbially precipitated carbonate boulders, bacterial mats, vesicomyid clam beds, frenulate and ampharetid beds, vestimentiferan tubeworm clumps, and fields of Bathysiphon filiformis tubes. Macrofaunal abundance increased and mean faunal delta C-13 signatures decreased in multicorer samples nearer the seep, suggesting that chemosynthetic production enhanced animal densities outside the seep center. Polychaetes dominated sediments, and ampharetids became especially abundant near microbial mats, while gastropods, hydroids, and sponges dominated carbonate rocks. A wide range of stable isotopic signatures reflected the diversity of microhabitats, and methane-derived carbon was the most prevalent source of nutrition for several taxa, especially those associated with carbonates. Megafaunal species living near the seep included longspine thornyhead (Sebastolobus altivelis), Pacific dover sole (Microstomus pacificus), and lithodid crabs (Paralomis verrilli), which represent targets for demersal fisheries. Sebastolobus altivelis was especially abundant (6.5-8.2 fish.100 m(-2)) and appeared to aggregate near the most active seep microhabitats. The Del Mar Methane Seep, like many others along the world's continental margins, exhibits diverse ecosystem functions and enhances regional diversity. Seeps such as this one may also contribute ecosystem services if they provide habitat for fishery species, export production to support margin food webs, and serve as sinks for methane-derived carbon.

Carbonate communities: The activity of anaerobic methane oxidizing microbes facilitates precipitation of vast quantities of authigenic carbonate at methane seeps. Here we demonstrate the significant role of carbonate rocks in promoting diversity by providing unique habitat and food resources for macrofaunal assemblages at seeps on the Costa Rica margin (400-1850 m). The attendant fauna is surprisingly similar to that in rocky intertidal shores, with numerous grazing gastropods (limpets and snails) as dominant taxa. However, the community feeds upon seep-associated microbes. Macrofaunal density, composition, and diversity on carbonates vary as a function of seepage activity, biogenic habitat and location. The macrofaunal community of carbonates at non-seeping (inactive) sites is strongly related to the hydrography (depth, temperature, O2) of overlying water, whereas the fauna at sites of active seepage is not. Densities are highest on active rocks from tubeworm bushes and mussel beds, particularly at the Mound 12 location (1000 m). Species diversity is higher on rocks exposed to active seepage, with multiple species of gastropods and polychaetes dominant, while crustaceans, cnidarians, and ophiuroids were better represented on rocks at inactive sites. Macro-infauna (larger than 0.3 mm) from tube cores taken in nearby seep sediments at comparable depths exhibited densities similar to those on carbonate rocks, but had lower diversity and different taxonomic composition. Seep sediments had higher densities of ampharetid, dorvilleid, hesionid, cirratulid and lacydoniid polychaetes, whereas carbonates had more gastropods, as well as syllid, chrysopetalid and polynoid polychaetes. Stable isotope signatures and metrics: The stable isotope signatures of carbonates were heterogeneous, as were the food sources and nutrition used by the animals. Carbonate δ13Cinorg values (mean = -26.98‰) ranged from -53.3‰ to +10.0‰, and were significantly heavier than carbonate δ13Corg (mean = -33.83‰), which ranged from -74.4‰ to -20.6‰. Invertebrates on carbonates had average δ13C (per rock) = -31.0‰ (range -18.5‰ to -46.5‰) and δ15N = 5.7‰ (range -4.5‰ to +13.4‰). Average δ13C values did not differ between active and inactive sites; carbonate fauna from both settings depend on chemosynthesis-based nutrition. Community metrics reflecting trophic diversity (SEAc, total Hull Area, ranges of δ13C and δ15N) and species packing (mean distance to centroid, nearest neighbor distance) also did not vary as a function of seepage activity or site. However, distinct isotopic signatures were observed among related, co-occurring species of gastropods and polychaetes, reflecting intense microbial resource partitioning. Overall, the substrate and nutritional heterogeneity introduced by authigenic seep carbonates act to promote diverse, uniquely adapted assemblages, even after seepage ceases. The macrofauna in these ecosystems remain largely overlooked in most surveys, but are major contributors to biodiversity of chemosynthetic ecosystems and the deep sea in general.

ABSTRACT Marine methane seeps are globally distributed geologic features in which reduced fluids, including methane, are advected upward from the subsurface. As a result of alkalinity generation during sulfate-coupled methane oxidation, authigenic carbonates form slabs, nodules, and extensive pavements. These carbonates shape the landscape within methane seeps, persist long after methane flux is diminished, and in some cases are incorporated into the geologic record. In this study, microbial assemblages from 134 native and experimental samples across 5,500 km, representing a range of habitat substrates (carbonate nodules and slabs, sediment, bottom water, and wood) and seepage conditions (active and low activity), were analyzed to address two fundamental questions of seep microbial ecology: (i) whether carbonates host distinct microbial assemblages and (ii) how sensitive microbial assemblages are to habitat substrate type and temporal shifts in methane seepage flux. Through massively parallel 16S rRNA gene sequencing and statistical analysis, native carbonates are shown to be reservoirs of distinct and highly diverse seep microbial assemblages. Unique coupled transplantation and colonization experiments on the seafloor demonstrated that carbonate-associated microbial assemblages are resilient to seep quiescence and reactive to seep activation over 13 months. Various rates of response to simulated seep quiescence and activation are observed among similar phylogenies (e.g., Chloroflexi operational taxonomic units) and similar metabolisms (e.g., putative S oxidizers), demonstrating the wide range of microbial sensitivity to changes in seepage flux. These results imply that carbonates do not passively record a time-integrated history of seep microorganisms but rather host distinct, diverse, and dynamic microbial assemblages. IMPORTANCE Since their discovery in 1984, the global distribution and importance of marine methane seeps have become increasingly clear. Much of our understanding of methane seep microorganisms—from metabolisms to community ecology—has stemmed from detailed studies of seep sediments. However, it has become apparent that carbonates represent a volumetrically significant habitat substrate at methane seeps. Through combined in situ characterization and incubation experiments, this study demonstrates that carbonates host microbial assemblages distinct from and more diverse than those of other seep habitats. This emphasizes the importance of seep carbonates as biodiversity locales. Furthermore, we demonstrate that carbonate-associated microbial assemblages are well adapted to withstand fluctuations in methane seepage, and we gain novel insight into particular taxa that are responsive (or recalcitrant) to changes in seep conditions.

The importance of tectonics and fluid flow in controlling cold seep habitats has long been appreciated at convergent margins but remains poorly understood in strike-slip systems. Here we present geophysical, geochemical, and biological data from an active methane seep offshore from Del Mar, California, in the inner California borderlands (ICB). The location of this seep appears controlled by localized transpression associated with a step in the San Diego Trough fault zone and provides an opportunity to examine the interplay between fluid expulsion and restraining step overs along strike-slip fault systems. These segment boundaries may have important controls on seep locations in the ICB and other margins characterized by strike-slip faulting (e.g., Greece, Sea of Marmara, and Caribbean). The strike-slip fault systems offshore southern California appear to have a limited distribution of seep sites compared to a wider distribution at convergent plate boundaries, which may influence seep habitat diversity and connectivity.

Mussels ( Mytilus edulis ) build massive, spatially complex, biogenic structures that alter the biotic and abiotic environment and provide a variety of ecosystem services. Unlike rocky shores, where mussels can attach to the primary substrate, soft sediments are unsuitable for mussel attachment. We used a simple lattice model, field sampling, and field and laboratory experiments to examine facilitation of recruitment (i.e., preferential larval, juvenile, and adult attachment to mussel biogenic structure) and its role in the development of power‐law spatial patterns observed in Maine, USA, soft‐bottom mussel beds. The model demonstrated that recruitment facilitation produces power‐law spatial structure similar to that in natural beds. Field results provided strong evidence for facilitation of recruitment to other mussels—they do not simply map onto a hard‐substrate template of gravel and shell hash. Mussels were spatially decoupled from non‐mussel hard substrates to which they can potentially recruit. Recent larval recruits were positively correlated with adult mussels, but not with other hard substrates. Mussels made byssal thread attachments to other mussels in much higher proportions than to other hard substrates. In a field experiment, mussel recruitment was highest to live mussels, followed by mussel shell hash and gravel, with almost no recruitment to muddy sand. In a laboratory experiment, evenly dispersed mussels rapidly self‐organized into power‐law clusters similar to those observed in nature. Collectively, the results indicate that facilitation of recruitment to existing mussels plays a major role in soft‐bottom spatial pattern development. The interaction between large‐scale resource availability (hard substrate) and local‐scale recruitment facilitation may be responsible for creating complex power‐law spatial structure in soft‐bottom mussel beds.

The deep sea is Earth's largest habitat but little is known about the nature of deep-sea parasitism. In contrast to a few characterized cases of bacterial and protistan parasites, the existence and biological significance of deep-sea parasitic fungi is yet to be understood. Here we report the discovery of a fungus-related parasitic microsporidium, Nematocenator marisprofundi n. gen. n. sp. that infects benthic nematodes at methane seeps on the Pacific Ocean floor. This infection is species-specific and has been temporally and spatially stable over 2 years of sampling, indicating an ecologically consistent host-parasite interaction. A high distribution of spores in the reproductive tracts of infected males and females and their absence from host nematodes' intestines suggests a sexual transmission strategy in contrast to the fecal-oral transmission of most microsporidia. N. marisprofundi targets the host's body wall muscles causing cell lysis, and in severe infection even muscle filament degradation. Phylogenetic analyses placed N. marisprofundi in a novel and basal clade not closely related to any described microsporidia clade, suggesting either that microsporidia-nematode parasitism occurred early in microsporidia evolution or that host specialization occurred late in an ancient deep-sea microsporidian lineage. Our findings reveal that methane seeps support complex ecosystems involving interkingdom interactions between bacteria, nematodes, and parasitic fungi and that microsporidia parasitism exists also in the deep-sea biosphere.

Ongoing greenhouse gas emissions can modify climate processes and induce shifts in ocean temperature, pH, oxygen concentration, and productivity, which in turn could alter biological and social systems. Here, we provide a synoptic global assessment of the simultaneous changes in future ocean biogeochemical variables over marine biota and their broader implications for people. We analyzed modern Earth System Models forced by greenhouse gas concentration pathways until 2100 and showed that the entire world's ocean surface will be simultaneously impacted by varying intensities of ocean warming, acidification, oxygen depletion, or shortfalls in productivity. In contrast, only a small fraction of the world's ocean surface, mostly in polar regions, will experience increased oxygenation and productivity, while almost nowhere will there be ocean cooling or pH elevation. We compiled the global distribution of 32 marine habitats and biodiversity hotspots and found that they would all experience simultaneous exposure to changes in multiple biogeochemical variables. This superposition highlights the high risk for synergistic ecosystem responses, the suite of physiological adaptations needed to cope with future climate change, and the potential for reorganization of global biodiversity patterns. If co-occurring biogeochemical changes influence the delivery of ocean goods and services, then they could also have a considerable effect on human welfare. Approximately 470 to 870 million of the poorest people in the world rely heavily on the ocean for food, jobs, and revenues and live in countries that will be most affected by simultaneous changes in ocean biogeochemistry. These results highlight the high risk of degradation of marine ecosystems and associated human hardship expected in a future following current trends in anthropogenic greenhouse gas emissions.

Upon their initial discovery, hydrothermal vents and methane seeps were considered to be related but distinct ecosystems, with different distributions, geomorphology, temperatures, geochemical properties and mostly different species. However, subsequently discovered vents and seep systems have blurred this distinction. Here, we report on a composite, hydrothermal seep ecosystem at a subducting seamount on the convergent Costa Rica margin that represents an intermediate between vent and seep ecosystems. Diffuse flow of shimmering, warm fluids with high methane concentrations supports a mixture of microbes, animal species, assemblages and trophic pathways with vent and seep affinities. Their coexistence reinforces the continuity of reducing environments and exemplifies a setting conducive to interactive evolution of vent and seep biota.

Mussels are important ecosystem engineers that have significant impacts on the ecology of the seafloor. In this study, we attempt to characterize and explain infaunal and epifaunal abundance patterns across scales in a spatially complex, intertidal, soft-bottom habitat: beds of the semi-infaunal mussel, Mytilus edulis. We used univariate and multivariate analyses in a nested sampling design at three locations in eastern Maine to show the relationships between environmental and macrofaunal assemblage variables. The results support the hypotheses that: 1) environmental and macrofaunal assemblage variables vary widely across spatial scales and exhibit maximum variance at different spatial scales; 2) abundance and diversity patterns of macrofauna are linked to spatial variation in environmental variables in the form of biogenic structure (live mussels and mussel shell hash), terrestrially-derived gravel, and, to a lesser degree, the proportion of silt-clay in the sediment; and 3) epifauna are generally positively correlated with hard-substrate features such as live mussels and gravel, and negatively correlated with the silt-clay content of the sediment, although the latter relationships were not strong. On the other hand, the results did not support the hypotheses that: 4) most infauna, specifically those with free-swimming larvae, are negatively correlated with live mussels and silt-clay proportion; instead, they were often positively correlated with live mussels and had weak, insignificant correlations with silt-clay; and 5) infauna without free-swimming larvae are positively correlated with live mussels and silt-clay proportion; instead, they showed some positive correlations with live mussels, but a mix of weak, insignificant correlations with silt-clay. Thus, mussel beds had a variety of scale-dependent relationships, both positive and negative, with each of the macrofaunal groups. The results show that biogenic structure in the form of live mussels and mussel shell hash is important in this system, as is terrestrial gravel, and that these variables are more important than silt-clay content for the macrofaunal assemblage. Mussel beds play an important role in regulating macrofaunal diversity in soft-bottom habitats, and their effects vary across spatial scales.