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Baited landers were deployed at 83 stations at four locations in the west Pacific Ocean from bathyal to hadal depths: The Kermadec Trench, the New Hebrides Trench, the adjoining South Fiji Basin and the Mariana Trench. Forty-seven putative fish species were observed. Distinct fish faunal groups were identified based on maximum numbers and percentage of observations. Both analyses broadly agreed on the community structure: A bathyal group at The absence of the abyssal group from the New Hebrides Trench and South Fiji Basin was due to the absence of macrourids ( Coryphaenoides spp.), which defined the group. The macrourids may be energetically limited in these areas. In their absence the species of the AHTZ group appear released of competition with the macrourids and are found far shallower at these sites. The fish groups had distinct feeding strategies while attending the bait: The bathyal and abyssal groups were almost exclusively necrophagous, the AHTZ group comprised predatory and generalist feeders, while the hadal snailfishes were exclusively predators. With increasing depth, predation was found to increase while scavenging decreased. The data suggest scavenging fish fauna do not extend deeper than the hadal boundary.

The snailfishes, family Liparidae (Scorpaeniformes), have found notable success in the hadal zone from ~6000−8200 m, comprising the dominant ichthyofauna in at least five trenches worldwide. Little is known about the biology of these deepest-living fishes, nor the factors that drive their success at hadal depths. Using recent collections from the Mariana Trench, Kermadec Trench, and neighboring abyssal plains, this study investigates the potential role of trophic ecology in structuring fish communities at the abyssal-hadal boundary. Stomach contents were analyzed from two species of hadal snailfishes, Notoliparis kermadecensis and a newly-discovered species from the Mariana Trench. Amphipods comprised the majority (Kermadec: 95.2%, Mariana: 97.4% index of relative importance) of stomach contents in both species. Decapod crustaceans, polychaetes ( N. kermadecensis only), and remains of carrion (squid and fish) were minor dietary components. Diet analyses of abyssal species (families Macrouridae, Ophidiidae, Zoarcidae) collected from near the trenches and the literature are compared to those of the hadal liparids. Stomachs from abyssal fishes also contained amphipods, however macrourids had a higher trophic plasticity with a greater diversity of prey items, including larger proportions of carrion and fish remains; supporting previous findings. Suction-feeding predatory fishes like hadal liparids may find an advantage to descending into the trench – where amphipods are abundant. More generalist feeders and scavengers relying on carrion, such as macrourids, might not benefit from this nutritional advantage at hadal depths. Compound specific isotope analysis of amino acids was used to estimate trophic level of these species (5.3±0.2 Coryphaenoides armatus , 5.2±0.2 C. yaquinae , 4.6±0.2 Spectrunculus grandis , 4.2±0.2 N. kermadecensis , 4.4±0.2 Mariana snailfish ) . Source amino acid δ 15 N values were especially high in hadal liparids (8.0±0.3‰ Kermadec, 6.7±0.2‰ Mariana), suggesting a less surface-derived food source than seen in the scavenging abyssal macrourids, C. armatus (3.5±0.3‰) and C. yaquinae (2.2±0.3‰). These results are compared to bulk muscle tissue isotopic compositions. This study provides the first comprehensive examination of the feeding ecology of the ocean's deepest-living fishes and informs new understanding of trophic interactions and fish community structure in and near the hadal zone.

Deep-sea fishes inhabit ∼75% of the biosphere and are a critical part of deep-sea food webs. Diet analysis and more recent trophic biomarker approaches, such as stable isotopes and fatty-acid profiles, have enabled the description of feeding guilds and an increased recognition of the vertical connectivity in food webs in a whole-water-column sense, including benthic-pelagic coupling. Ecosystem modeling requires data on feeding rates; the available estimates indicate that deep-sea fishes have lower per-individual feeding rates than coastal and epipelagic fishes, but the overall predation impact may be high. A limited number of studies have measured the vertical flux of carbon by mesopelagic fishes, which appears to be substantial. Anthropogenic activities are altering deep-sea ecosystems and their services, which are mediated by trophic interactions. We also summarize outstanding data gaps.

As agencies shift from single-species management to ecosystem-based fisheries management, ecosystem models are gaining interest for understanding species dynamics in relation to oceanographic and ecological processes and human marine uses. However, information on community structure or distribution of many species that occupy deep (>30 m) waters is largely unavailable. We amassed a total of 24 686 fish observations of 523 species/taxa for the 30−410 m depth areas surrounding the main Hawaiian Islands (MHI). We also obtained estimates of geomorphological variables, including substrate type, slope, rugosity, and ridge-like features. Using these 2 data sources, we (1) identified distinct fish communities along the 30−410 m depth gradient, and (2) generated relative biomass maps for fish functional groups. We showed that the mesophotic zone ranges between 30 and 129 m, with a fish faunal break at 60 m. Beyond this zone, 4 subphotic zones were identified: upper rariphotic (130−169 m), mid-rariphotic (170−239 m), lower rariphotic (240−319 m), and upper bathyal (320−410 m). We assigned fish species to functional groups partly based on identified depth ranges and fitted general additive models (GAMs) integrating geomorphological covariates to the functional group relative biomass estimates to determine the environmental variables that best predict the probability of encounter and relative biomass of each fish functional group. Finally, GAM predictions were employed to map functional group relative biomass distributions. These distribution maps showed a high relative biomass of many groups in the center of the MHI chain. This study contributes to a better understanding of fish community structure around the MHI and will inform ecosystem model parameterization.

Hadal ocean sediments, found at sites deeper than 6,000m water depth, are thought to contain microbial communities distinct from those at shallower depths due to high hydrostatic pressures and higher abundances of organic matter. These communities may also differ from one other as a result of geographical isolation. Here we compare microbial community composition in surficial sediments of two hadal environments-the Mariana and Kermadec trenches-to evaluate microbial biogeography at hadal depths. Sediment microbial consortia were distinct between trenches, with higher relative sequence abundances of taxa previously correlated with organic matter degradation present in the Kermadec Trench. In contrast, the Mariana Trench, and deeper sediments in both trenches, were enriched in taxa predicted to break down recalcitrant material and contained other uncharacterized lineages. At the 97% similarity level, sequence-abundant taxa were not trench-specific and were related to those found in other hadal and abyssal habitats, indicating potential connectivity between geographically isolated sediments. Despite the diversity of microorganisms identified using culture-independent techniques, most isolates obtained under in situ pressures were related to previously identified piezophiles. Members related to these same taxa also became dominant community members when native sediments were incubated under static, long-term, unamended high-pressure conditions. Our results support the hypothesis that there is connectivity between sediment microbial populations inhabiting the Mariana and Kermadec trenches while showing that both whole communities and specific microbial lineages vary between trench of collection and sediment horizon depth. This in situ biodiversity is largely missed when incubating samples within pressure vessels and highlights the need for revised protocols for high-pressure incubations.

The International Seabed Authority (ISA) is developing regulations to control the future exploitation of deep-sea mineral resources including sulphide deposits near hydrothermal vents, polymetallic nodules on the abyssal seafloor, and cobalt crusts on seamounts. Under the UN Convention on the Law of the Sea the ISA is required to adopt are taking measures to ensure the effective protection of the marine environment from harmful effects arising from mining-related activities. Contractors are required to generate environmental baselines and assess the potential environmental consequences of deep seafloor mining. Understandably, nearly all environmental research has focused on the seafloor where the most direct mining effects will occur. However, sediment plumes and other impacts (e.g., noise) from seafloor mining are likely to be extensive in the water column. Sediment plumes created on the seafloor will affect the benthic boundary layer which extends 10s to 100s of meters above the seafloor. Separation or dewatering of ore from sediment and seawater aboard ships will require discharge of a dewatering plume at some depth in the water column. It is important to consider the potential impacts of mining on the ocean’s midwaters (depths from ~200 m to the seafloor) because they provide vital ecosystem services and harbor substantial biodiversity. The better known epipelagic or sunlit surface ocean provisions the rest of the water column through primary production and export flux (This was not the focus at this workshop as the subject was considered too large and surface discharges are unlikely). It is also home to a diverse community of organisms including commercially important fishes such as tunas, billfish, and cephalopods that contribute to the economies of many countries. The mesopelagic or twilight zone (200-1000 m) is dimly lit and home to very diverse and abundant communities of organisms. Mesopelagic plankton and small nekton form the forage base for many deep-diving marine mammals and commercially harvested epipelagic species. Furthermore, detritus from the epipelagic zone falls through the mesopelagic where it is either recycled, providing the vital process of nutrient regeneration, or sinks to greater depths sequestering carbon from short-term atmospheric cycles. The waters below the mesopelagic down to the seafloor (both the bathypelagic and abyssopelagic) are very poorly characterized but are likely large reservoirs of novel biodiversity and link the surface and benthic ecosystems. Great strides have been made in understanding the biodiversity and ecosystem function of the ocean’s midwaters, but large regions, including those containing many exploration license areas and the greater depths where mining plumes will occur, remain very poorly studied. It is clear that pelagic communities are distinct from those on the seafloor and in the benthic boundary layer. They are often sampled with different instrumentation. The fauna have relatively large biogeographic ranges and they are more apt to mix freely across stakeholder boundaries, reference areas and other spatial management zones. Pelagic organisms live in a three-dimensional habitat and their food webs and populations are vertically connected by daily or lifetime migrations and the sinking flux of detritus from the epipelagic. The fauna do not normally encounter hard surfaces, making them fragile, and difficult to capture and maintain for sensitivity or toxicity studies. Despite some existing general knowledge, ecological baselines for midwater communities and ecosystems that likely will be impacted by mining have not been documented. There is an urgent need to conduct more research and evaluate the midwater biota (microbes to fishes) in regions where mining is likely to occur. Deep-sea mining activities may affect midwater organisms in a number of ways, but it is still unclear at what scale perturbations may occur. The sediment plumes both from collectors on the seafloor and from midwater discharge will have a host of negative consequences. They may cause respiratory distress from clogged gills or respiratory surfaces. Suspension feeders, such as copepods, polychaetes, salps, and appendicularians, that filter small particles from the water and form an important basal group of the food web, may suffer from dilution of their food by inorganic sediments and/or clogging of their fragile mucous filter nets. Small particles may settle on gelatinous plankton causing buoyancy issues. Metals, including toxic elements that will enter the food web, will be released from pore waters and crushed ore materials. Sediment plumes will also absorb light and change backscatter properties, reducing visual communication and bioluminescent signaling that are very important for prey capture and reproduction in midwater animals. Noise from mining activities may alter the behaviors of marine mammals and other animals. Small particles have high surface area to volume ratios, high pelagic persistence and dispersal and as a result greater potential to result in pelagic impacts. All of these potential effects will result in mortality, migration (both horizontal and vertical), decreased fitness, and shifts in community composition. Depending on the scale and duration of these effects, there could be reduction in provisioning to commercial fish species, delivery of toxic metals to pelagic food webs and hence human seafood supply, and alterations to carbon transport and nutrient regeneration services. After four days of presentations and discussions, the workshop participants came to several conclusions and synthesized recommendations. 1. Assuming no discharge in the epipelagic zone, it is essential to minimize mining effects in the mesopelagic zone because of links to our human seafood supply as well as other ecosystem services provided by the mesopelagic fauna. This minimization could be accomplished by delivering dewatering discharge well below the mesopelagic/bathypelagic transition (below ~1000 m depth). 2. Research should be promoted by the ISA and other bodies to study the bathypelagic and abyssopelagic zones (from ~1000 m depths to just above the seafloor). It is likely that both collector plumes and dewatering plumes will be created in the bathypelagic, yet this zone is extremely understudied and contains major unknowns for evaluating mining impacts. 3. Management objectives, regulations and management actions need to prevent the creation of a persistent regional scale “haze” (enhanced suspended particle concentrations) in pelagic midwaters. Such a haze would very likely cause chronic harm to deep midwater ecosystem biodiversity, structure and function. 4. Effort is needed to craft suitable standards, thresholds, and indicators of harmful environmental effects that are appropriate to pelagic ecosystems. In particular, suspension feeders are very important ecologically and are likely to be very sensitive to sediment plumes. They are a high priority for study. 5. Particularly noisy mining activities such as ore grinding at seamounts and hydrothermal vents is of concern to deep diving marine mammals and other species. One way to minimize sound impacts would be to minimize activities in the sound-fixing-and-ranging (SOFAR) channel (typically at depths of ~1000 m) which transmits sounds over very long distances. 6. A Lagrangian (drifting) perspective is needed in monitoring and management because the pelagic ecosystem is not a fixed habitat and mining effects are likely to cross spatial management boundaries. For example, potential broad-scale impacts to pelagic ecosystems should be considered in the deliberations over preservation reference zones, the choice of stations for environmental baseline and monitoring studies and other area-based management and conservation measures. 7. Much more modeling and empirical study of realistic mining sediment plumes is needed. Plume models will help evaluate the spatial and temporal extent of pelagic (as well as benthic) ecosystem effects and help to assess risks from different technologies and mining scenarios. Plume modeling should include realistic mining scenarios (including duration) and assess the spatial-temporal scales over which particle concentrations exceed baseline levels and interfere with light transmission to elucidate potential stresses on communities and ecosystem services. Models should include both near and far field-phases, incorporating realistic near field parameters of plume generation, flocculation, particle sinking, and other processes. It is important to note that some inputs to these models such as physical oceanographic parameters are lacking and should be acquired in the near-term. Plume models need to be complemented by studies to understand effects on biological components by certain particle sizes and concentrations.

Hadal environments sustain diverse microorganisms. A few studies have investigated hadal microbial communities consisting of free-living or particle-associated bacteria and archaea. However, animal-associated microbial communities in hadal environments remain largely unexplored, and comparative analyses of animal gut microbiota between two isolated hadal environments have never been done so far. In the present study, 228 Gb of gut metagenomes of the giant amphipod Hirondellea gigas from two hadal trenches, the Mariana Trench and Japan Trench, were sequenced and analyzed. Taxonomic analysis identified 49 microbial genera commonly shared by the gut microbiota of the two H. gigas populations. However, the results of statistical analysis, in congruency with the alpha and beta diversity analyses, revealed significant differences in gut microbial composition across the two trenches. Abundance variation of Psychromonas, Propionibacterium, and Pseudoalteromonas species was observed. Microbial cooccurrence was demonstrated for microbes that were overrepresented in the Mariana trench. Comparison of functional potential showed that the percentage of carbohydrate metabolic genes among the total microbial genes was significantly higher in the guts of H. gigas specimens from the Mariana Trench. Integrating carbon input information and geological characters of the two hadal trenches, we propose that the differences in the community structure might be due to several selective factors, such as environmental variations and microbial interactions. IMPORTANCE The taxonomic composition and functional potential of animal gut microbiota in deep-sea environments remain largely unknown. Here, by performing comparative metagenomics, we suggest that the gut microbial compositions of two Hirondellea gigas populations from the Mariana Trench and the Japan Trench have undergone significant divergence. Through analyses of functional potentials and microbe-microbe correlations, our findings shed light on the contributions of animal gut microbiota to host adaptation to hadal environments.

The effective evaluation of trophic interactions in pelagic food webs is essential for understanding food web ecology, conservation biology, and management. We tested the applicability of compound-specific isotope analysis of amino acids (CSIA-AA) for (1) characterizing trophic positions (TPs) of nine species from four trophic groups (tunas, squids, myctophids, and euphausiids) within a pelagic food web in the eastern tropical Pacific (ETP) Ocean, (2) evaluating trophic discrimination factors (TDFs) of each trophic group, and (3) detecting spatial changes in TPs and food chain length across a region with heterogeneous productivity. Although δ15N values of bulk tissues generally increased from south-to-north, CSIA-AA revealed that trophic positions were uniform throughout our study area. These results suggest that variability in δ15N values were largely driven by nitrogen cycling dynamics in the ETP, which highlights the importance of these processes for the interpretation of δ15N values in food web studies. Absolute TP estimates were unrealistic for higher-level species, and TDFs (tunas: 4.0‰, squids: 4.6‰, myctophids: 5.0‰, and euphausiids: 7.0‰) were lower than a widely used ecosystem TDF. We used remotely sensed oceanographic data to evaluate the physical oceanography and biological productivity throughout our study area and found significant relationships between δ15N values, nitrate concentrations, and SST across our study area. We did not find a gradient in phytoplankton cell size co-occurring with an expected productivity gradient across our sampling region, which substantiated our isotope results indicating non-significant spatial changes in TP and food chain length across the ETP.

The aim of this paper is to measure metabolic rates of megafauna living in depths greater than 6000 m. Echinoderms, actinarians and a polychaete were captured by remotely operated vehicle (ROV) and inserted into respiration chambers in situ at depths of 4049 m, 7140 m and 8074 m in the region of the Kermadec Trench SW Pacific Ocean. Hadal research has moved into a new frontier as technological improvements now allow for a meticulous investigation of trench ecology in depths greater than 6000 m. The development of an in situ respirometer for use in these studies was deployed in the Kermadec Trench to obtain the first ever rates of basal metabolic rates of hadal megafauna. Typical deep-sea experiments of individual animal physiology must deal with covarying factors of pressure, temperature, light and food supply in this study investigated the effects of pressure and increased food supply on overall animal metabolism. In the Kermadec Trench, holothurian respiration rates (n=4), 0.079±0.011 (mean±SE) µmol-O 2 g −1 h −1 , were higher than those captured at abyssal depths (n=2), 0.018±0.002 µmol-O 2 g -1 h −1 , in the same region (p 10 adjusted to a common temperature of 2.5 °C trench holothurian respiration rates ranged between 0.068 and 0.119 µmol-O 2 g −1 h −1 . Anemone respiration rates were remarkably similar between abyssal and hadal specimens, 0.110 and 0.111 µmol-O 2 g −1 h −1 , respectively. Our results on echinoderm respiration when corrected for temperature and mass fall below the slope regression when compared with other in situ measurements at shallower ocean depths.

A major limitation to fully integrated ecosystem based fishery management approaches is a lack of information on the spatial distribution of marine species and the environmental conditions shaping these distributions. This is particularly problematic for deep-water species that are hard to sample and are data poor. The past decade has seen the rapid development of a suite of advanced species distribution, or ecological niche, modelling approaches developed specifically to support efficient and targeted management. However, model performance can vary significantly and the appropriateness of which methods are best for a given application remains questionable. Species distribution models were developed for three commercially valuable Hawaiian deep-water eteline snappers: Etelis coruscans (Onaga), Etelis carbunculus (Ehu) and Pristipomoides filamentosus (Opakapaka). Distributional data for these species was relatively sparse. To identify the best method, model performance and distributional accuracy was assessed and compared using three approaches: Generalised Additive Models (GAM), Boosted Regression Trees (BRT) and Maximum Entropy (MaxEnt). Independent spatial validation data found MaxEnt consistently provided better model performance with ‘good’ model predictions (AUC =>0.8). Each species was influenced by a unique combination of environmental conditions, with depth, terrain (slope) and substrate (low lying unconsolidated sediments), being the three most important in shaping their distributions. Sustainable fisheries management, marine spatial planning and environmental decision support systems rely on an understanding species distribution patterns and habitat linkages. This study demonstrates that predictive species distribution modelling approaches can be used to accurately model and map sparse species distribution data across marine landscapes. The approach used herein was found to be an accurate tool to delineate species distributions and associated habitat linkages, account for species-specific differences and support sustainable ecosystem-based management.

Baited stereo-camera surveys of fish assemblages provide conservative estimates of abundance and length-frequency distributions. While underwater camera systems have numerous advantages over traditional fishing and diver surveys, limitations in sampling capacity, data processing time, and resultant data still exist. Previous studies have shown that shorter camera soak times can increase sampling efficiency and reduce per-sample data processing time without affecting overall data quality. Using data from stereo-video surveys of bottomfish in the main Hawaiian Islands, this study evaluates the effect of camera soak time on relative abundance metrics, fish length data, sampling efficiency, and power to detect differences in relative abundance and fish lengths. A soak time of 15 min was found to be the shortest duration able to capture bottomfish abundance and length metrics while 30 min generated data that did not significantly differ from the standard 40-min soak time. These shorter soak times allow for better survey efficiency and improved cost–benefit through increased levels of field sampling and reductions in video-processing time, while maintaining the power to detect differences in bottomfish relative abundance and lengths. The main drawback to shortening soak time was the concurrent reduction in the number of length measurements collected per species. An increased sample yield can alleviate this effect but only for bottomfish with a higher frequency of occurrence. Species-specific patterns in abundance were apparent in this study suggesting a strong influence of fish behavior on stereo-video abundance metrics. While a soak time of 15 to 30 min was found to be sufficient for effectively sampling bottomfish, the cost–benefit of employing a given soak time in future stereo-video surveys should be assessed based on the target species and survey goals.

Phytoplankton production drives marine ecosystem trophic-structure and global fisheries yields. Phytoplankton biomass is particularly influential near coral reef islands and atolls that span the oligotrophic tropical oceans. The paradoxical enhancement in phytoplankton near an island-reef ecosystem—Island Mass Effect (IME)—was first documented 60 years ago, yet much remains unknown about the prevalence and drivers of this ecologically important phenomenon. Here we provide the first basin-scale investigation of IME. We show that IME is a near-ubiquitous feature among a majority (91%) of coral reef ecosystems surveyed, creating near-island ‘hotspots' of phytoplankton biomass throughout the upper water column. Variations in IME strength are governed by geomorphic type (atoll vs island), bathymetric slope, reef area and local human impacts (for example, human-derived nutrient input). These ocean oases increase nearshore phytoplankton biomass by up to 86% over oceanic conditions, providing basal energetic resources to higher trophic levels that support subsistence-based human populations., The Island Mass Effect (IME), where island proximity enhances phytoplankton biomass, remains a poorly understood phenomenon. Here, the authors show the IME is a common feature among Pacific reefs, driving increased production that creates biological hotspots in otherwise barren ocean basins.

The photochemical degradation of the neurotoxin methylmercury (MeHg) in marine surface waters is of great interest because it reduces the amount of MeHg available for uptake and bioaccumulation in marine aquatic food webs. Studies have shown that the dominant cause of odd isotope mass independent fractionation (odd-MIF) of Hg in marine foodwebs is the photo-degradation of MeHg. Residual MeHg is then incorporated into low trophic level organisms and bio-accumulated without additional mass independent fractionation. Based on this understanding of Hg isotope fractionation we sought to use Hg isotope measurements of fish tissues containing Hg mostly as MeHg to assess the relative degree of photochemical decomposition across the world's oceans. In 19 samples of flying fish, the magnitude of odd-MIF varies by a factor of ~2. We estimate that 56 to 80% of MeHg was photo-degraded prior to entering the food web depending on location. The proportion of MeHg degradation does not correlate with latitude, solar radiation or estimates of the concentration of DOC or chlorophyll at the collection sites, but it does correlate with proxies for water clarity. The ratio of odd-MIF for 199Hg compared to 201Hg is constant in all flying fish sampled (1.20 ± 0.03) suggesting that there is a common mechanism for photo-degradation of MeHg in surface waters across oceans and hemispheres. The ratio of odd-MIF to mass dependent fractionation (MDF) is generally consistent with photochemical degradation but is more variable (2.76 ± 0.33), suggesting that there is negligible internal demethylation of MeHg in flying fish and that variable amounts of MDF occur at different locations in the ocean, likely driven by variable rates of methylation and demethylation of Hg in the water column.

Midwater zooplankton are major agents of biogeochemical transformation in the open ocean; however their characteristics and activity remain poorly known. Here we evaluate midwater zooplankton biomass, amino acid (AA)-specific stable isotope composition (δ15N values) using compound-specific isotope analysis of amino acids (CSIA-AA), trophic position, and elemental composition in the North Pacific Subtropical Gyre (NPSG). We focus on zooplankton collected in the winter, spring, and summer to evaluate midwater trophic dynamics over a seasonal cycle. For the first time we find that midwater zooplankton respond strongly to seasonal changes in production and export in the NPSG. In summer, when export from the euphotic zone is elevated and this ‘summer pulse’ material is transported rapidly to depth, CSIA-AA indicates that large particles (>53 μm) dominate the food web base for zooplankton throughout the midwaters, and to a large extent even into the upper bathypelagic zone. In winter, when export is low, zooplankton in the mid-mesopelagic zone continue to rely on large particle basal resources, but resident zooplankton in the lower mesopelagic and upper bathypelagic zones switch to include smaller particles (0.7–53 μm) in their food web base, or even a subset of the small particle pool. Midwater zooplankton migration patterns also vary with season, with migrants distributed more evenly at night through the euphotic zone in summer as compared to being more compressed in the upper mixed layer in winter. Deeper zooplankton migration within the mesopelagic zone is also reduced in late summer, likely due to the increased magnitude of large particle material available at depth during this season. Our observed seasonal change in activity and trophic dynamics drives modestly greater biomass in summer than winter through the mesopelagic zone. In contrast midwater zooplankton carbon (C), nitrogen (N), and phosphorus (P) composition does not change with season. Instead we find increasing C:N, C:P, and N:P ratios with greater depths, likely due to decreases in proteinaceous structures and organic P compounds and increases in storage lipids with depth. Our study highlights the importance and diversity of feeding strategies for small zooplankton in NPSG midwaters. Many small zooplankton, such as oncaeid and oithonid copepods, are able to access small particle resources at depth and may be an important trophic link between the microbial loop and deep dwelling micronekton species that also rely on small particle-based food webs. Our observed midwater zooplankton trophic response to export-driven variation in the particle field at depth has important implications for midwater metabolism and the export of C to the deep sea.

The demersal ichthyofauna at the abyssal Station M at 4100 m depth in the NE Pacific Ocean (34°50′N, 123° 00′W) has been sampled from 1989 onwards using a combination of techniques; baited traps, long lines, otter trawls, towed camera sledge, remotely operated vehicles (ROV), human occupied vehicle (Alvin), benthic long-term time lapse camera and baited camera landers. The fish fauna is dominated by two species of macrourid Coryphaenoides armatus and C. yaquinae. Other species present are Bathysaurus mollis (Bathysauridae), Coryphaenoides leptolepis (Macrouridae), Bassozetus nasus, Spectrunculus grandis (Ophidiidae), three species of Liparidae, including Genioliparis ferox and a Zoarcid (Pachycara sp.). Evidence of deposition of egg masses by snail fishes, liparids on oceanographic equipment is discussed.

As a unique but poorly investigated habitat within marine ecosystems, hadal trenches have received interest in recent years. This study explores the gut microbial composition and function in hadal amphipods, which are among the dominant carrion feeders in hadal habitats. Further analyses of a dominant strain revealed genomic features that may contribute to its adaptation to the amphipod gut environment. Our findings provide new insights into animal-associated bacteria in the hadal biosphere., Amphipods are the dominant scavenging metazoan species in the Mariana Trench, the deepest known point in Earth’s oceans. Here the gut microbiota of the amphipod Hirondellea gigas collected from the Challenger and Sirena Deeps of the Mariana Trench were investigated. The 11 amphipod individuals included for analyses were dominated by Psychromonas, of which a nearly complete genome was successfully recovered (designated CDP1). Compared with previously reported free-living Psychromonas strains, CDP1 has a highly reduced genome. Genome alignment showed deletion of the trimethylamine N-oxide (TMAO) reducing gene cluster in CDP1, suggesting that the “piezolyte” function of TMAO is more important than its function in respiration, which may lead to TMAO accumulation. In terms of nutrient utilization, the bacterium retains its central carbohydrate metabolism but lacks most of the extended carbohydrate utilization pathways, suggesting the confinement of Psychromonas to the host gut and sequestration from more variable environmental conditions. Moreover, CDP1 contains a complete formate hydrogenlyase complex, which might be involved in energy production. The genomic analyses imply that CDP1 may have developed adaptive strategies for a lifestyle within the gut of the hadal amphipod H. gigas. IMPORTANCE As a unique but poorly investigated habitat within marine ecosystems, hadal trenches have received interest in recent years. This study explores the gut microbial composition and function in hadal amphipods, which are among the dominant carrion feeders in hadal habitats. Further analyses of a dominant strain revealed genomic features that may contribute to its adaptation to the amphipod gut environment. Our findings provide new insights into animal-associated bacteria in the hadal biosphere.

The present study examined metabolic enzyme activities of 61 species of demersal fishes (331 individuals) trawled from a 3000 m depth range. Citrate synthase, lactate dehydrogenase, malate dehydrogenase, and pyruvate kinase activities were measured as proxies for aerobic and anaerobic activity and metabolic rate. Fishes were classified according to locomotory mode, either benthic or benthopelagic. Fishes with these two locomotory modes were found to exhibit differences in metabolic enzyme activity. This was particularly clear in the overall activity of citrate synthase, which had higher activity in benthopelagic fishes. Confirming earlier, less comprehensive studies, enzyme activities declined with depth in benthopelagic fishes. For the first time, patterns in benthic species could be explored and these fishes also exhibited depth-related declines in enzyme activity, contrary to expectations of the visual interactions hypothesis. Trends were significant when using depth parameters taken from the literature as well as from the present trawl information, suggesting a robust pattern regardless of the depth metric used. Potential explanations for the depth trends are discussed, but clearly metabolic rate does not vary simply as a function of mass and habitat temperature in fishes as shown by the substantial depth-related changes in enzymatic activities.

We used bulk tissue d 13 C and d 15 N values and d 15 N values of individual amino acids (AA) to characterize the trophic structure of a pelagic fish assemblage from the North Pacific Subtropical Gyre (NPSG) ecosystem. We focus on energy flow between fishes inhabiting distinct epipelagic, mesopelagic, and upper bathypelagic habitats and on predatory fish foraging across and within these depth habitats. Trophic positions (TPs) estimated from a combination of trophic and source AA d 15 N values (TPTr-Src) spanned a narrow range of 0.7 TP for 10 species of large fishes, including tunas, billfishes, and gempylids (TPTr-Src 4.3-5.0). Similarly, 13 species of small micronekton fishes encompassed a range of 1.2 TP (TPTr-Src 2.6-3.8). The d 15 N values of three source AAs were found to increase with increasing depth of capture across the 13 micronekton fish species (d 15 NPhe range 56.6&; d 15 NGly range 513.4&; d 15 NSer range 513.6&), indicating that some species from epipelagic, mesopelagic, and upper bathypelagic communities access distinct food resources, such as suspended particles. These isotopic depth trends are consistent with previous observations in particulate organic matter and zooplankton from the NPSG, providing new evidence that large pelagic and micronekton fishes access a food web fueled by particles formed in surface waters but that are highly modified by microbes as they slowly settle to remote depths. On the contrary, no significant relationships between the d 15 N values of source AAs and habitat depth were observed in the large predator fish group, of which many species move and forage across large depth gradients.

We assess the metabolic demand of mesopelagic zooplankton for carbon, nitrogen, and phosphorus in the North Pacific Subtropical Gyre (NPSG). We compare zooplankton metabolic demand with the attenuation of sinking particle fluxes, and find the average metabolic demand for resident midwater zooplankton can account for 1.3× the loss of sinking particulate organic carbon and particulate nitrogen fluxes, and 2.6× the loss of particulate phosphorus fluxes. Zooplankton metabolic demand for carbon remains significant (0.4-1.9×) relative to the loss in sinking particulate fluxes, even when new depth-specific dry weight conversion factors and recent global-bathymetric models of zooplankton metabolism are applied. These new models reduce zooplankton carbon demands to reasonably match particle flux attenuation in the mesopelagic zone. Zooplankton metabolic demand for phosphorus is particularly large in comparison to particle flux attenuation (1.5-9×), and when temporal change in the molar carbon : phosphorus ratio of the attenuation in particulate fluxes are considered, mesopelagic zooplankton in the NPSG may episodically become phosphorus-limited. Midwater zooplankton have the potential to be important mediators of carbon flux to the deep ocean in the NPSG.

The complex biogeochemical cycle of Hg makes identifying primary sources of fish tissue Hg problematic. To identify sources and provide insight into this cycle, we combined carbon (δ13C), nitrogen amino acid (δ15NPhe), and Hg isotope (Δ199Hg, Δ201Hg, δ202Hg) data for six species of Hawaiian marine bottomfish. Results from these isotopic systems identified individuals within species that likely fed from separate food webs. Terrestrial freshwater inputs to coastal sediments were identified as the primary source of tissue Hg in the jack species, Caranx ignobilis, which inhabit shallow marine ecosystems. Thus, coastal C. ignobilis were a biological vector transporting Hg from freshwater environments into marine ecosystems. Depth profiles of Hg isotopic compositions for bottomfish (excludung C. ignobilis) were similar, but not identical, to profiles for open-ocean pelagic fishes, suggesting that in both settings inorganic Hg, which was ultimately transformed to monomethylmercury (MeHg) and bioaccumulated, was ...

There is growing interest in mining polymetallic nodules from the abyssal Clarion-Clipperton Zone (CCZ) in the tropical Pacific Ocean. Despite having been the focus of environmental studies for decades, the benthic megafauna of the CCZ remain poorly known. To predict and manage the environmental impacts of mining in the CCZ, baseline knowledge of the megafauna is essential. The ABYSSLINE Project has conducted benthic biological baseline surveys in the UK Seabed Resources Ltd polymetallic-nodule exploration contract area (UK-1). Prior to ABYSSLINE research cruises in 2013 and 2015, no biological studies had been done in this area of the eastern CCZ. Using a Remotely Operated Vehicle and Autonomous Underwater Vehicle (as well as several other pieces of equipment), the megafauna within the UK Seabed Resources Ltd exploration contract area (UK-1) and at a site ~250 km east of the UK-1 area were surveyed, allowing us to make the first estimates of megafaunal morphospecies richness from the imagery collected. Here, we present an atlas of the abyssal annelid, arthropod, bryozoan, chordate, ctenophore and molluscan megafauna observed and collected during the ABYSSLINE cruises to the UK-1 polymetallic-nodule exploration contract area in the CCZ. There appear to be at least 55 distinct morphospecies (8 Annelida, 12 Arthropoda, 4 Bryozoa, 22 Chordata, 5 Ctenophora, and 4 Mollusca) identified mostly by morphology but also using molecular barcoding for a limited number of animals that were collected. This atlas will aid the synthesis of megafaunal presence/absence data collected by contractors, scientists and other stakeholders undertaking work in the CCZ, ultimately helping to decipher the biogeography of the megafauna in this threatened habitat.

We have developed a global biogeographic classification of the mesopelagic zone to reflect the regional scales over which the ocean interior varies in terms of biodiversity and function. An integrated approach was necessary, as global gaps in information and variable sampling methods preclude strictly statistical approaches. A panel combining expertise in oceanography, geospatial mapping, and deep-sea biology convened to collate expert opinion on the distributional patterns of pelagic fauna relative to environmental proxies (temperature, salinity, and dissolved oxygen at mesopelagic depths). An iterative Delphi Method integrating additional biological and physical data was used to classify biogeographic ecoregions and to identify the location of ecoregion boundaries or inter-regions gradients. We define 33 global mesopelagic ecoregions. Of these, 20 are oceanic while 13 are ‘distant neritic.’ While each is driven by a complex of controlling factors, the putative primary driver of each ecoregion was identified. While work remains to be done to produce a comprehensive and robust mesopelagic biogeography (i.e., reflecting temporal variation), we believe that the classification set forth in this study will prove to be a useful and timely input to policy planning and management for conservation of deep-pelagic marine resources. In particular, it gives an indication of the spatial scale at which faunal communities are expected to be broadly similar in composition, and hence can inform application of ecosystem-based management approaches, marine spatial planning and the distribution and spacing of networks of representative protected areas. publishedVersion

Many deep-sea fishes have a gelatinous layer, or subdermal extracellular matrix, below the skin or around the spine. We document the distribution of gelatinous tissues across fish families (approx. 200 species in ten orders), then review and investigate their composition and function. Gelatinous tissues from nine species were analysed for water content (96.53 ± 1.78% s.d.), ionic composition, osmolality, protein (0.39 ± 0.23%), lipid (0.69 ± 0.56%) and carbohydrate (0.61 ± 0.28%). Results suggest that gelatinous tissues are mostly extracellular fluid, which may allow animals to grow inexpensively. Further, almost all gelatinous tissues floated in cold seawater, thus their lower density than seawater may contribute to buoyancy in some species. We also propose a new hypothesis: gelatinous tissues, which are inexpensive to grow, may sometimes be a method to increase swimming efficiency by fairing the transition from trunk to tail. Such a layer is particularly prominent in hadal snailfishes (Liparidae); therefore, a robotic snailfish model was designed and constructed to analyse the influence of gelatinous tissues on locomotory performance. The model swam faster with a watery layer, representing gelatinous tissue, around the tail than without. Results suggest that the tissues may, in addition to providing buoyancy and low-cost growth, aid deep-sea fish locomotion.

Paraliparis hawaiiensis n.sp. is described from the north-western Hawaiian Islands from two specimens collected at 2196 and 3055 m. It differs from other North Pacific Ocean species in its chin pore arrangement, tooth pattern and body proportions. Although liparid specimens have previously been collected from Hawaii, they were undescribed and are now lost. Therefore, this is the first liparid species described from the archipelago. In situ photographs of Hawaiian snailfishes are also shown and discussed here.

Ecosystem-based fisheries management strategies require knowledge of trophic relationships. Trophic position (TP) estimates from compound specific nitrogen isotopic analysis of amino acids (AA-CSIA) show promise as the method can disentangle confounding factors associated with changing δ15N values at the base of the food web, but it has yet to be tested in many organisms. This novel technique requires two empirically determined biological parameters: 1) β, the difference in δ15N values between glutamic acid (glu) and phenylalanine (phe) in primary producers and 2) trophic enrichment factor (TEF), the 15N enrichment of glu and phe at each trophic step. Values of β (3.4‰) and TEF (7.6‰) have been suggested for animals in aquatic environments; however recent observations indicate that TEF valuesmay be variable, particularly among elasmobranchs where urea retentionmay alter nitrogen isotope fractionation between glu and phe. To test these uncertainties,we determined TEF values for three species of sharks, sand tiger (Carcharias taurus), lemon (Negaprion brevirostris), and leopard sharks (Triakis semifasciata), and one teleost species, opakapaka (Pristipomoides filamentosus) grown on controlled and well characterized diets for durations ranging from three (T. semifasciata) to over five years (P. filamentosus). TEF values for both elasmobranchs and opakapaka were ~2‰, significantly lower than TEFs previously reported. These results do not support the hypothesis that urea retention lowers 15N trophic enrichment between glu and phe in elasmobranchs. Rather, isotopic enrichment factors may be primarily driven by differences in dietary protein quality, leading to distinct TEFs for herbivores (~7.6‰) and carnivores (b7.6‰). We propose a method to calculate TP which integrates different TEF values for herbivores and carnivores.

No fish have been found in the deepest 25% of the ocean (8,400-11,000 m). This apparent absence has been attributed to hydrostatic pressure, although direct evidence is wanting because of the lack of deepest-living species to study. The common osmolyte trimethylamine N-oxide (TMAO) stabilizes proteins against pressure and increases with depth, going from 40 to 261 mmol/kg in teleost fishes from 0 to 4,850 m. TMAO accumulation with depth results in increasing internal osmolality (typically 350 mOsmol/kg in shallow species compared with seawater's 1,100 mOsmol/kg). Preliminary extrapolation of osmolalities of predicted isosmotic state at 8,000-8,500 m may indicate a possible physiological limit, as greater depths would require reversal of osmotic gradients and, thus, osmoregulatory systems. We tested this prediction by capturing five of the second-deepest known fish, the hadal snailfish (Notoliparis kermadecensis; Liparidae), from 7,000 m in the Kermadec Trench. We found their muscles to have a TMAO content of 386 ± 18 mmol/kg and osmolality of 991 ± 22 mOsmol/kg. These data fit previous extrapolations and, combined with new osmolalities from bathyal and abyssal fishes, predict isosmotic state at 8,200 m. This is previously unidentified evidence that biochemistry could constrain the depth of a large, complex taxonomic group.

Particle dynamics are an essential component of global ocean biogeochemistry as they transport essential nutrients, carbon, and other reactive elements and compounds from the surface ocean to depth in the water column. The North Pacific Ocean is characterized by spatial and temporal variations in particle export mediated by a diverse food web and variations in environmental conditions, such as oxygenation. Here we explored temporal variability in the downward flux of particulate carbon (PC) and nitrogen (PN) using 238U 234Th disequilibria, sediment traps and in situ pumps in winter, spring and summer at the time-series site Station ALOHA, a region characterized by a well-defined summer export pulse that influences the composition and structure of the biological community within the mesopelagic zone. We further explored spatial variability in PC and PN fluxes along a latitudinal gradient (17.5°-5°N x 150°W) that transitions from a low to high productivity region influenced by equatorial upwelling, with several stations further characterized by a shallow (~ 130 m) oxygen minimum zone. Winter PC and PN fluxes at 150 m at Station ALOHA were low, while summer and spring fluxes were significantly higher, coinciding with a seasonal export pulse associated with diazotrophs. PC and PN fluxes along the 155°W transect were also low at 150 m and similar to those measured at Station ALOHA in winter. At Station ALOHA zooplankton impart a greater influence over both small and large PC (and PN) fluxes in February relative to September or May, when heterotrophic bacteria play a proportionally larger role in particle remineralization and cycling. Along the transect stations, PC fluxes were too low to discern any clear trends with latitude, likely due to El Nino conditions at the time of sampling. Where vertical water column profiles of PC and PN fluxes were available, PC (and PN) fluxes were found to peak in the subsurface at 8°N (50 m) and 5°N (75 m) with zooplankton grazing and microbial remineralization following patterns similar to those found in February at Station ALOHA. Combined, these results support the hypothesis that small and large particles contribute to mesopelagic carbon demand depending on season, with smaller particles having greater contributions to mesopelagic food webs when surface derived particle export is low.

Owing to the paucity of data on the red muscle of deep-sea fishes, the goal of this study was to determine the proportions of red muscle in demersal fishes and its enzymatic activities to characterize how routine swimming abilities change with depths of occurrence. Cross sectional analysis of the trunk musculature was used to evaluate the proportion of red muscle in 38 species of Californian demersal fishes living at depths between 100 and 3000 m. The activity of metabolic enzymes was also assayed in a sub-set of 18 species. Benthic fishes had lower proportions of red muscle and lower metabolic enzyme activities than benthopelagic species. Mean proportion of red muscle declined significantly with depth with the greatest range of values in shallow waters and species with low proportions found at all depths. This suggested that while sedentary species occur at all depths, the most active species occur in shallow waters. Citrate synthase activity declined significantly with depth across all species, indicating that the mass-specific metabolic capacity of red muscle is lower in deep-sea species. These patterns may be explained by coupling of red and white muscle physiologies, a decrease in physical energy of the environment with depth or by the prevalence of anguilliform body forms and swimming modes in deep-living species.

The diets of 5 large predatory mesopelagic fishes—Alepisaurus ferox (longnosed lancetfish), Gempylus serpens (snake mackerel), Lepidocybium flavobrunneum (Smith's escolar), and Lampris spp. (big-eye and small-eye opah, or moonfish)—from the central North Pacific Ocean (around Hawaii) were examined (n = 430, all species combined), most for the first time. Recent analysis of fishery data has shown that many of these species have been undergoing decadal increases in abundance, suggesting system-wide changes. A. ferox diet was numerically dominated by hyperiid amphipods from 3 genera (Phrosina, Phronima, Platyscelus; 37%N), pelagic polychaete worms, mesopelagic fishes (including young A. ferox size classes), and cephalopods. G. serpens fed primarily on epipelagic fishes (exocoetids, molids) and ommastrephid squids. Diets of the 2 Lampris species were the most similar to one another, consisting of large numbers and frequent occurrences of the onychoteuthid squid Walvisteuthis youngorum and a diverse assemblage of epipelagic and mesopelagic fishes. More than 90% of the L. flavobruneum stomachs were without food items; small numbers of prey identified included the ommastrephid squid Sthenoteuthis oualaniensis, aristeid shrimps, and unidentified fishes. The diet descriptions support the idea that these predatory fishes carve out unique ecological niches in the pelagic envi- ronment by exploiting unique components of micronekton communities across epipelagic and mesopelagic depth zones. Adult size classes of tunas and billfishes occupying a shared vertical habitat do not appear to compete for prey resources to any great extent, perhaps allowing for suc- cessful partitioning of limited prey resources within an oligotrophic gyre ecosystem.

The success of marine protected areas (MPAs) as a tool for conservation and fisheries management has been well documented. However, these results have typically been seen in shallow water systems and questions remain whether this management strategy could be successfully applied to deepwater ecosystems. Our objectives were to determine the efficacy of four deepwater MPAs called bottomfish restricted fishing areas (BRFAs), with various time spans of protection, monitored at depths between 90 and 310 m from 2007 to 2011 for six species of deepwater snapper and one grouper harvested in the Main Hawaiian Islands. Our results suggested that the duration of protection influenced reserve effects, particularly for target species. Mean fish length, and in some cases abundance, increased for one or more of the most economically important target species inside nearly all tested BRFAs. In addition, more mature fish were seen inside the BRFA with the longest duration of protection (~14 years); species richness increased outside this area while inside it remained the same. Here, we provide the first evidence that deepwater MPAs can have positive effects on deepwater species and that many protection effects were consistent with results found in shallow water ecosystems. While these findings are novel, additional data over greater temporal scales will be necessary to determine whether these trends will continue and if others will become important over time.

We used amino acid (AA) compound-specific isotope analysis (d15NAA and d13CAA values) of midwater zooplankton and suspended particles to examine their dynamics in the mesopelagic zone. Suspended particle d15NAA values increased by up to 14% with depth, whereas particle trophic status (measured as trophic position, TP) remained constant at 1.6 6 0.07. Applying a Rayleigh distillation model to these results gave an observed kinetic isotope fractionation of 5.7 6 0.4%, similar to that previously measured for protein hydrolysis. AA-based degradation index values also decreased with depth on the particles, whereas a measure of heterotrophic resynthesis (SV) remained constant at 1.2 6 0.3. The main mechanism driving 15N enrichment of suspended particles appears to be isotope fractionation associated with heterotrophic degradation, rather than a change in trophic status or N source with depth. In zooplankton the ‘‘source’’ AA phenylalanine (Phe) became 15N enriched by up to 3.5% with depth, whereas zooplankton TP increased by up to 0.65 between the surface ocean and midwaters. Both changes in the d15N values of food resources at the base of the zooplankton food web and changes in zooplankton TP drive observed zooplankton 15N enrichment with depth. Midwater zooplankton d15NPhe values were lower by 5–8% compared with suspended particles, indicating this organic matter pool is not a significant zooplankton food resource at depth. Instead, 62–88% of the N sustaining midwater zooplankton is surface derived, obtained through consumption of sinking particles, carnivory of vertical migrants, or direct feeding in surface waters at night.

We document a systematic decline in 202 Hg, 1 199 Hg and 1 201 Hg values with the depth at which fish feed. We show that these mercury isotope trends can be explained only if monomethylmercury is produced below the surface mixed layer, including in the underlying oxygen minimum zone, that is, between 50 and more than 400 m depth. Specifically, we estimate that about 20‐40% of the monomethylmercury detected below the surface mixed layer originates from the surface and enters deeper waters either attached to sinking particles, or in zooplankton and micronekton that migrate to depth. We suggest that the remaining monomethylmercury found at depth is produced below the surface mixed layer by methylating microbes that live on sinking particles. We suggest that microbial production of monomethylmercury below the surface mixed later contributes significantly to anthropogenic mercury uptake into marine food webs.

Consumption rates of marine predators are vital to assessing their trophic impacts and potential consequences of fisheries removal and habitat alteration, yet are rarely estimated. Standard metabolic rates were estimated for juvenile brown stingrays, Dasyatis lata, and used as input parameters for a bioenergetics model to predict consumption rates. Temperature and mass had significant effects on metabolic rates. The energy budget of juvenile brown stingrays was heavily weighted toward metabolism, accounting for 66 % of consumed energy. Growth accounted for 7 % of the energy budget indicating very slow growth potentially due to limited food resources. Population consumption rates suggest potential for strong top-down effects on prey populations due to stingray predation. This study suggests the use of Kāne‘ohe Bay as a nursery habitat for juvenile brown stingrays is a trade-off between increased juvenile survival through predator avoidance and a late age at first maturity due to slow growth rates resulting from low prey availability.

The effects of low-pH, high-pCO2 conditions on deep-sea organisms were examined during four deep-sea CO2 release experiments simulating deep-ocean C sequestration by the direct injection of CO2 into the deep sea. We examined the survival of common deep-sea, benthic organisms (microbes; macrofauna, dominated by Polychaeta, Nematoda, Crustacea, Mollusca; megafauna, Echinodermata, Mollusca, Pisces) exposed to low-pH waters emanating as a dissolution plume from pools of liquid carbon dioxide released on the seabed during four abyssal CO2-release experiments. Microbial abundance in deep-sea sediments was unchanged in one experiment, but increased under environmental hypercapnia during another, where the microbial assemblage may have benefited indirectly from the negative impact of low-pH conditions on other taxa. Lower abyssal metazoans exhibited low survival rates near CO2 pools. No urchins or holothurians survived during 30–42 days of exposure to episodic, but severe environmental hypercapnia during one experiment (E1; pH reduced by as much as ca. 1.4 units). These large pH reductions also caused 75% mortality for the deep-sea amphipod, Haploops lodo, near CO2 pools. Survival under smaller pH reductions (ΔpHo0.4 units) in other experiments (E2, E3, E5) was higher for all taxa, including echinoderms. Gastropods, cephalopods, and fish were more tolerant than most other taxa. The gastropod Retimohnia sp. and octopus Benthoctopus sp. survived exposure to pH reductions that episodically reached −0.3 pH units. Ninety percent of abyssal zoarcids (Pachycara bulbiceps) survived exposure to pH changes reaching ca. −0.3 pH units during 30–42 day-long experiments.

There have been numerous reports of plastic debris accumulation in surface waters of the central North Pacific Subtropical Gyre. Further, incidences have been reported of plastic ingestion by different marine organisms, including seabirds and small planktivorous fishes. Diet studies (2007 to 2012) of predatory pelagic fishes from this general region showed repeat observa- tions of anthropogenic marine debris ingestion in 7 species (n = 595 individuals examined). Inci- dence rates ranged from

Little is known about the metabolism of deep-living, benthic invertebrates, despite its importance in estimating energy flow through individuals and populations. To evaluate the effects of depth and broad taxonomic group/locomotory mode, we measured the respiration rates of 25 species of benthic decapod crustaceans and 18 species of echinoderms from the littoral zone to the deep slope of Hawaii. Specimens were collected by hand, trap, or submersible and maintained in the laboratory at temperatures close to ambient temperatures recorded at the time of collection. After acclimatization to laboratory conditions, oxygen consumption was measured for each individual in closed chambers. Overall, crustaceans had higher metabolic rates than echinoderms, and within the crustaceans, caridean shrimps had higher rates than crabs and lobsters. These differences are probably related to locomotory mode and general levels of activity. At in situ environmental temperatures, metabolic rates of deeper-living invertebrates are much lower than those of shallower living species, but this decline is explained by changes in temperature. When the data were compared with similar data sets collected off California and in the Mediterranean, Hawaiian crabs, lobsters, and echinoderms had lower metabolic rates than similar species in the other regions after adjustments for temperature were made. Some of these differences could be methodological. Regional food web models should use broad taxonomic groupings and region-specific data when possible.

This study provides the first comprehensive fishery-independent baseline assessment of commercially important deepwater bottomfish populations across the main Hawaiian Islands. Differences in bottomfish relative abundance and size distribution were evaluated for 6 deepwater Bottomfish Restricted Fishing Areas (BRFAs). While no differences were detected in species relative abundance, evaluation of size-frequency distributions found the 2 most commercially valuable species (Etelis coruscans and Pristipomoides filamentosus) to be significantly larger inside the BRFA at Ni‘ihau, located off the most remote of the main Hawaiian Islands. This BRFA is 1 of 2 ongoing BRFAs offering 10 yr of protection. This result highlighted the time it may take a long-lived and slow-growing species to show a detectable response to protection and that size distribution analyses can detect these more subtle changes. No positive effects of protection were detected for the second ongoing BRFA located off Hawai‘i. Instead, 2 species (P. filamentosus and P. sieboldii) were significantly larger outside the BRFA. In contrast to Ni‘ihau, the second BRFA established in 1998 originally included less preferred habitat and is next to the second largest port in Hawai‘i, offering greater access, higher population pressure and more problematic enforcement. This study demonstrates that biological, sociological and environmental context must also be considered when interpreting the effectiveness of marine protected areas.

Deep-sea communities rely on epipelagic surface production as a primary source of energy and food. The flux of phytodetritus drives many abyssal ecological processes but the flux of large particles such as nekton carcasses, macroalgae, and wood may also be important. Recent baited camera experiments noted that some abyssal fish consumed spinach and phytoplankton placed on the seafloor. To evaluate if fish or other scavengers would consume natural plant or macroalgal material falling to the deep-sea floor we conducted camera experiments using Sargassum or mackerel bait in the Sargasso Sea. A benthic community of invertebrates was attracted to Sargassum, which naturally falls to the seafloor in this area. In five instances it was observed that an isopod Bathyopsurus sp. removed a piece of Sargassum from the main clump and left the field of view with it. An ophiuroid is also observed handling a piece of Sargassum. The group of scavengers attracted to mackerel bait was very different and was dominated by large ophidiid fish. In contrast to studies elsewhere in the abyssal North Atlantic, only a small number of rattails are observed, which could be related to water depth or an ichthyofaunal zonal change between oligotrophic and eutrophic regions.

This paper presents results demonstrating detection of cardiac motion and heart rate monitoring by radar for fish using a contact antenna, as a less-invasive alternative to surgical electrocardiogram (ECG) measurements. Tilapia are used as test subjects with a patch antenna. The frequency of operation, 2.4 GHz, is chosen as a tradeoff between antenna size and propagation. Single lead ECG measurements are used as a reference for comparison with two electrode configurations. The heart rate detected by the radar matched the heart rate detected by ECG. With water temperature near 30°C, measured heart rates varied with sedation level and over time, in the range of 30-80 bpm.

Here we consider the role of depth as a driver of evolution in a genus of deep-sea fishes. We provide a phylogeny for the genus Coryphaenoides (Gadiformes: Macrouridae) that represents the breadth of habitat use and distributions for these species. In our consensus phylogeny species found at abyssal depths (4000m) form a well-supported lineage, which interestingly also includes two non-abyssal species, C. striaturus and C. murrayi, diverging from the basal node of that lineage. Biogeographic analyses suggest the genus may have originated in the Southern and Pacific Oceans where contemporary species diversity is highest. The abyssal lineage seems to have arisen secondarily and likely originated in the Southern/Pacific Oceans but diversification of this lineage occurred in the Northern Atlantic Ocean. All abyssal species are found in the North Atlantic with the exception of C. yaquinae in the North Pacific and C. filicauda in the Southern Ocean. Abyssal species tend to have broad depth ranges and wide distributions, indicating that the stability of the deep oceans and the ability to live across wide depths may promote population connectivity and facilitate large ranges. We also confirm that morphologically defined subgenera do not agree with our phylogeny and that the Giant grenadier (formerly Albatrossia pectoralis) belongs to Coryphaenoides, indicating that a taxonomic revision of the genus is needed. We discuss the implications of our findings for understanding the radiation and diversification of this genus, and the likely role of adaptation to the abyss.

Climate change is expected to impact all aspects of marine ecosystems, including fisheries. Here, we use output from a suite of 11 earth system models to examine projected changes in two ecosystem-defining variables: temperature and food availability. In particular, we examine projected changes in epipelagic temperature and, as a proxy for food availability, zooplankton density. We find that under RCP8.5, a high business-as-usual greenhouse gas scenario, increasing temperatures may alter the spatial distribution of tuna and billfish species richness across the North Pacific basin. Furthermore, warmer waters and declining zooplankton densities may act together to lower carrying capacity for commercially valuable fish by 2-5% per decade over the 21st century. These changes have the potential to significantly impact the magnitude, composition, and distribution of commercial fish catch across the pelagic North Pacific. Such changes will in turn ultimately impact commercial fisheries' economic value. Fishery managers should anticipate these climate impacts to ensure sustainable fishery yields and livelihoods.

12 páginas, 6 figuras, 5 tablas., There is a general hypothesis that species inhabiting deep-sea waters have lower fecundity and larger eggs than shallower species. However, there are few comparative studies which explore this trend because of the complexity of sampling in deep waters, especially in fishes. We present here the first analysis of fecundity and egg size with depth along an isothermal environment. We calculate the relative fecundity and egg size of 11 species of demersal deep-sea fish from the western Mediterranean and included in our analyses published data for an additional 14 species from the same geographic area. The results show that the relative fecundity (eggs per g of individual) of the analyzed fishes slightly decreased along the bathymetric gradient, whereas egg size increased with depth. When the analysis was conducted including only species from the order Gadiformes, the most speciose group in the region and with the widest depth range of distribution (50–2000 m), there was no relationship between relative fecundity and depth, while the deepest species had larger egg sizes than shallower ones. The finding of similar relative fecundities but larger egg sizes suggests that these deep-sea species are investing a higher amount of energy in the production of offspring than shallower water counterparts. The results are discussed in relation to the isothermal characteristics of the deep Mediterranean Sea and ecological adaptations for reproductive success., This study was conducted within the framework oft he RECS(REN02/04556/C02/MAR), PROMETEO(CTM2007-66316-C02/MAR) and DOSMARES (CTM2010-21810-C03-03/MAR) projects by the Spanish Science and Innovation Ministry respectively, to JBC.

The ability of six grenadier species from the North Atlantic, North Pacific, and Southern Ocean to sustain deep-sea fisheries is assessed. These species are captured in high amounts as bycatch and a few are taken in targeted fisheries, yet population status for most is poorly known or known for only a small portion of their range. A productivity and susceptibility analysis showed that none of the species was highly productive, which was not unexpected given their life history characteristics. While grenadiers were ranked more vulnerable than species in the northeastern Pacific groundfish fisheries, none of the investigated species was ranked as highly susceptible or heavily exploited. This result exposed several weaknesses in the PSA technique and attribute scoring. Management actions and regulations are discussed, which, if employed, might make grenadier fisheries sustainable., Sí

Metabolic enzyme activities and gill surface areas were measured across 10 species of demersal fishes from Monterey Canyon, California, which features a prominent oxygen minimum zone (OMZ). Comparisons were made between species living both within and outside of the OMZ. Enzyme activities showed no significant trend toward aerobic suppression or heightened reliance on anaerobic metabolism in response to the OMZ. While flatfish species living both within and outside of the OMZ had similarly low enzyme activities, the OMZ-dwelling Microstomus pacificus had 1.8–3 times larger gill surface area than comparably sized flatfishes from higher-oxygen waters, suggesting a morphological adaptation to low oxygen. In scorpaeniform fishes, high aerobic metabolism was accompanied by large gill surface areas in two routine-swimming OMZ-dwelling species (Anoplopoma fimbria and Careproctus melanurus). Low aerobic activities and small gills were found in two Sebastolobus species, suggesting a low oxygen demand resulting from a more sedentary behavior compared to other Scorpaeniformes. In gadiform fishes, no differences were measured in enzyme activity levels, but larger gill surface areas were measured in OMZdwelling Nezumia liolepis. These results indicate adaptation to low oxygen in a variety of ways that balance oxygen demand with supply, with no indication that these species rely on enhanced anaerobic metabolism. With both flatfishes and rattails, adaptation to OMZs is demonstrated through increased gill surface area. Oxygen minimum zones (OMZs) are midwater regions (200–1000 m) where dissolved oxygen levels are reduced by an order of magnitude relative to waters above and below the OMZ core, defined by a concentration of . 0.5 mL O2

Biochemical indices of white (WM) and red muscle (RM) aerobic and anaerobic metabolic capacity were measured in 14 species of benthic and benthopelagic chondrichthyans from a depth of ~90 to 2,200 m to evaluate the relationship between metabolic capacity and depth of occurrence, phylogeny, and locomotor mode. Maximal activities of the enzymes citrate synthase, malate dehydrogenase (MDH), lactate dehydrogenase (LDH), and pyruvate kinase (PK) were analyzed in muscle tissue at 10 °C. These were combined with previously published elasmobranch data in order to represent a comprehensive range of depths, phylogeny, and locomotor modes (i.e., benthic, benthopelagic, pelagic). Significant decreases in WM PK and LDH activities and a lack of significant trends in RM enzyme activities with increasing median depth of occurrence (MDO) indicate a depth-related reduction in both burst-locomotor and metabolic capacity. These trends are consistent with the “visual-interactions hypothesis.” Phylogeny and locomotor mode had little influence on enzyme activities compared to MDO, and the present study suggests similar activities in co-occurring demersal sharks and rays. Overall, the present study indicates low metabolic capacities in deep-sea chondrichthyans, which is important to consider when managing deep-sea fisheries.

Submarine canyons are reported to be sites of enhanced fish biomass and productivity on continental margins. However, little is known about the effects of canyons on fish biodiversity, in particular on oceanic islands, which are imbedded in regions of low productivity. Using submersibles and high- definition video surveys, we investigated demersal fish assemblages in two submarine canyons and slope areas off the island of Moloka'i, Hawai'i, at depths ranging from 314 to 1100 m. We addressed the interactions between the abundance, species richness and composition of the fish assemblage, and organic matter input and habitat heterogeneity, testing the hypotheses that heterogeneous bottom habitats and higher organic matter input in canyons enhance demersal fish abundance, and species density, richness and diversity, thereby driving differences in assemblage structure between canyons and slopes. Sediment type, substrate inclination, water-mass properties (temperature and dissolved oxygen) and organic matter input (modeled POC flux and percent detritus occurrence) were put into multivariate multiple regression models to identify potential drivers of fish assemblage structure. A total of 824 fish were recorded during � 13 h of video yielding 55 putative species. Macrouridae was the most diverse family with 13 species, followed by Congridae (5), Ophidiidae (4) and Halosauridae (3). Assemblage structure changed markedly with depth, with the most abrupt change in species composition occurring between the shallowest stratum (314-480 m) and intermediate and deep strata (571-719 m, 946-1100 m). Chlorophthalmus sp. dominated the shallow stratum, macrourids and synaphobranchid eels at intermediate depths, and halosaurs in the deepest stratum. Assemblages only differed significantly between canyon and slope habitats for the shallow stratum, and the deep stratum at one site. Dissolved oxygen explained the greatest proportion of variance in the multivariate data, followed by POC flux and percent organic-detritus occurrence. Fish abundances were generally higher in canyons but only statistically significant for the deepest stratum. Reduced fish abundances both in canyon and slope transects occurred at intermediate depths within the core of the oxygen minimum zone (OMZ). Species density, diversity and richness and abundance were usually higher in the canyons, but only statistically higher in the deepest stratum. Possible causes for increased abundance and species densities and richness in the deepest stratum in canyons include reduced disturbance at deeper depths. We conclude that submarine canyons on oceanic islands are likely to be sites of enhanced fish abundance and species richness, but that these enhancing effects are offset when oxygen concentra- tions fall below � 0.7 ml l

Proximate composition, bulk tissue and amino acid compound-specific nitrogen isotopic analyses (CSIA) were used to determine whether dietary differences were responsible for the spatial variability in growth of spiny lobster and slipper lobster in the Northwestern Hawaiian Islands (NWHI). Abdominal tissues were collected and analyzed from both species at Necker Island and Maro Reef from 2006 to 2008. Protein and lipid levels did not differ significantly between locations in either species. Bulk tissue 15N of both species was significantly negatively correlated with growth for both species; however, the analysis assumed constant isotopic composition of autotrophs across this region. CSIA, which accounts for 15N variability at the base of the food web, indicated that spiny lobsters at both locations occupied the same trophic position whereas the slower-growing Maro Reef slipper lobsters fed at a lower trophic position relative to Necker Island slipper lobsters. Spatial variability in the abundance or diversity of preferred prey items appears to be responsible for the spatial variability in growth and the specific morphology and behavior of these species dictated how they coped with dietary restraints. These findings increase the understanding of NWHI coral reef ecosystem processes as well as highlight dangers of using consumer bulk tissue isotopic data without considering variation in the nitrogen isotopic composition at the base of the food web.