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Characterization of the biological, physical, and chemical properties of a toxic thin layer in a temperate marine system

The distribution of plankton in the ocean is patchy across a wide range of spatial and temporal scales. One type of oceanographic feature that exemplifies this patchiness is a ‘thin layer’. Thin layers are subsurface aggregations of plankton that range in vertical thickness from centimeters to a few meters, which may extend horizontally for kilometers and persist for days. We undertook a field campaign to characterize the biological, physical, and chemical properties of thin layers in Monterey Bay, California (USA), an area where these features can be persistent. The particle aggregates (marine snow) sampled in the study had several quantifiable properties indicating how the layer was formed and how its structure was maintained. Particles were more elongated above the layer, and then changed orientation angle and increased in size within the layer, suggesting passive accumulation of particles along a physical gradient. The shift in particle aggregate orientation angle near the pycnocline suggests that shear may also have played a role in generating the thin layer. Pseudo-nitzschia spp. were the most abundant phytoplankton within the thin layer. Further, both dissolved and particulate domoic acid were highest within the thin layer. We suggest that phosphate stress is responsible for the formation of Pseudo-nitzschia spp. aggregates. This stress together with increased nitrogen in the layer may lead to increased bloom toxicity in the subsurface blooms of Pseudo-nitzschia spp. Several zooplankton groups were observed to aggregate above and below the layer. With the knowledge that harmful algal bloom events can occur in subsurface thin layers, modified sampling methods to monitor for these hidden incubators could greatly improve the efficacy of early-warning systems designed to detect harmful algal blooms in coastal waters.

Marine Ecology Progress Series (MEPS)

Warming sea surface temperatures fuel summer epidemics of eelgrass wasting disease

Seawater temperatures are increasing, with many unquantified impacts on marine diseases. While prolonged temperature stress can accelerate host-pathogen interactions, the outcomes in nature are poorly quantified. We monitored eelgrass wasting disease (EWD) from 2013-2017 and correlated mid-summer prevalence of EWD with remotely sensed seawater temperature metrics before, during, and after the 2015-2016 marine heatwave in the northeast Pacific, the longest marine heatwave in recent history. Eelgrass shoot density declined by 60% between 2013 and 2015 and did not recover. EWD prevalence ranged from 5-70% in 2013 and increased to 60-90% by 2017. EWD severity approximately doubled each year between 2015 and 2017. EWD prevalence was positively correlated with warmer temperature for the month prior to sampling while EWD severity was negatively correlated with warming prior to sampling. This complex result may be mediated by leaf growth; bigger leaves may be more likely to be diseased, but may also grow faster than lesions, resulting in lower severity. Regional stressors leading to population declines prior to or early in the heatwave may have exacerbated the effects of warming on eelgrass disease susceptibility and reduced the resilience of this critical species.

Marine Ecology Progress Series

Dermal denticle shedding rates vary between two captive shark species

Shark dermal scale (denticle) accumulation in the fossil record can provide information about the abundance and composition of past shark communities. Denticles are shed continuously, such that a single shark leaves a scattered composite of many isolated denticles in sediments. However, the rate of denticle shedding as well as how these rates vary among shark species with different life modes and their consistency over time are unknown, limiting the interpretation of denticle assemblages. To better understand the process of denticle shedding and calibrate the relationship between absolute shark abundance in the environment and denticle deposition in sediments, we captured denticles shed by 2 shark species in a large aquarium over 9 mo. We then simulated how these aquarium-derived shedding rates shape the relationship between shark abundance and denticle accumulation. Bonnethead sharks Sphyrna tiburo , a more active, benthopelagic species with small, thin denticles, shed 3.6 times faster on average than zebra sharks Stegostoma fasciatum , a more sedentary, demersal species with large, robust denticles. This pattern persisted when shedding rates were corrected by estimated denticle quantities, shark space use, and methodological factors (2.2- to 3.8-fold difference). Over the study, bonnethead shark shedding rates declined while zebra shark shedding rates increased slightly. Finally, denticle assemblage composition corresponded with the relative abundance of denticles on the body of each species, consistent with natural shedding rather than selective loss. Overall, we show that shark taxa contribute unevenly to the denticle record, indicating that shedding rate measurements can help inform and constrain ecological interpretations of denticle assemblages.

Marine Ecology Progress Series

Phoretic sharksuckers (Echeneis naucrates) associated with an elasmobranch host occupy higher relative trophic positions

The relationship between phoretic diskfishes and their hosts is a classic example of marine symbiosis, yet surprisingly few studies have quantified this trophic relationship. We investigated the hypothesis that by consuming host parasites and prey scraps phoretic diskfishes (Echeneidae) feed at a higher relative trophic position than free-living individuals through expanded foraging opportunities. We used carbon and nitrogen stable isotope analysis of muscle tissue from both free-living and commensal sharksuckers Echeneis naucrates and their hosts, supplemented with gut-content analysis, to investigate this hypothesis. Our analysis revealed that commensal sharksuckers likely occupy higher relative trophic positions than free-living sharksuckers. The importance of scavenging host prey decreased ontogenetically as sharksuckers shifted from symbiotic phoresis to free-swimming behavior, leading to an ontogenetic change in which obligately commensal juveniles occupy higher trophic positions than free-living adults which are only facultatively commensal. Relative differences in δ 13 C and δ 15 N among individual host-commensal pairs varied among host species, suggesting potential differences in foraging opportunities among host taxa.

Marine Ecology Progress Series

Kinetic determinations of trace element bioaccumulation in the mussel Mytilus edulis

Laboratory experiments employing radiotracer methodology were conducted to determine the assimilation efficiencies from ingested natural seston, the influx rates from the dissolved phase and the efflux rates of 6 trace elements (Ag, Am, Cd, Co, Se and Zn) in the mussel Mytilus edulis. A kinetic model was then employed to predict trace element concentration in mussel tissues in 2 locations for which mussel and environmental data are well described: South San Francisco Bay (California, USA) and Long Island Sound (New York, USA). Assimilation efficiencies from natural seston ranged from 5 to 18% for Ag, 0.6 to 1% for Am, 8 to 20% for Cd, 12 to 16% for Co, 28 to 34% for Se, and 32 to 41% for Zn. Differences in chlorophyll a concentration in ingested natural seston did not have significant impact on the assimilation of Am, Co, Se and Zn. The influx rate of elements from the dissolved phase increased with the dissolved concentration, conforming to Freundlich adsorption isotherms. The calculated dissolved uptake rate constant was greatest for Ag, followed by Zn > Am = Cd > Co > Se. The estimated absorption efficiency from the dissolved phase was 1.53% for Ag, 0.34% for Am, 0.31% for Cd, 0.11% for Co, 0.03% for Se and 0.89% for Zn. Salinity had an inverse effect on the influx rate from the dissolved phase and dissolved organic carbon concentration had no significant effect on trace element uptake. The calculated efflux rate constants for all elements ranged from 1.0 to 3.0% d-1. The route of trace element uptake (food vs dissolved) and the duration of exposure to dissolved trace elements (12 h vs 6 d) did not significantly influence trace element efflux rates. A model which used the experimentally determined influx and efflux rates for each of the trace elements, following exposure from ingested food and from water, predicted concentrations of Ag, Cd, Se and Zn in mussels that were directly comparable to actual tissue concentrations independently measured in the 2 reference sites in national monitoring programs. Sensitivity analysis indicated that the total suspended solids load, which can affect mussel feeding activity, assimilation, and trace element concentration in the dissolved and particulate phases, can significantly influence metal bioaccumulation for particle-reactive elements such as Ag and Am. For all metals, concentrations in mussels are proportionately related to total metal load in the water column and their assimilation efficiency from ingested particles. Further, the model predicted that over 96% of Se in mussels is obtained from ingested food, under conditions typical of coastal waters. For Ag, Am, Cd, Co and Zn, the relative contribution from the dissolved phase decreases significantly with increasing trace element partition coefficients for suspended particles and the assimilation efficiency in mussels of ingested trace elements; values range between 33 and 67% for Ag, 5 and 17% for Am, 47 and 82% for Cd, 4 and 30% for Co, and 17 and 51% for Zn.

Marine Ecology Progress Series

Estimating Pacific walrus abundance and survival with multievent mark-recapture models

Arctic marine ecosystems are undergoing rapid physical and biological change associated with climate warming and loss of sea ice. Sea ice loss will impact many species through altered spatial and temporal availability of resources. In the Bering and Chukchi Seas, the Pacific walrus Odobenus rosmarus divergens is one species that could be impacted by rapid environmental change, and thus, population assessments are needed to monitor changes in the status of this ecologically and culturally important marine mammal. We conducted a 5 yr genetic mark-recapture study to estimate demographic parameters for the Pacific walrus. We developed a Bayesian multievent mark-recapture model to estimate walrus survival and abundance while accounting for age misclassification. We estimated the probability of juvenile annual survival as 0.63 (95% credible interval [CrI]: 0.39-0.87) and adult female annual survival as 0.90 (95% CrI: 0.74-1.00). We estimated total abundance as 257 193 (95% CrI: 171 138-366 366). We provide the first estimate of total Pacific walrus abundance since an aerial survey in 2006, which generated a substantially less precise total population size estimate (129 000; 95% CI: 55 000-507 000). The emerging ecosystem state in the northern Bering and Chukchi Seas will likely result in a decline in Pacific walrus abundance, but there is substantial uncertainty regarding the magnitude of the anticipated decline. Our demographic estimates provide critical information to evaluate future population trends of this subsistence resource vital to communities that border the Bering and Chukchi Seas in the USA and Russia.

Alaska

Laysan albatross exhibit complex behavioral plasticity in the subtropical and subarctic North Pacific Ocean

Animals that regularly traverse habitat extremes between the subtropics and subarctic are expected to exhibit foraging behaviors that respond to changes in dynamic ocean habitats, and these behaviors may facilitate adaptations to novel and changing climates. During the chick-provisioning stage, Laysan albatross Phoebastria immutabilis parents regularly undertake short- and long-distance foraging trips throughout the vast central North Pacific Ocean. We examined GPS tracking data among chick-provisioning albatrosses in Hawai‘i to characterize habitats during short- and long-distance trips. The study period encompassed a marine heatwave (2014) and the cooling period after an extreme El Niño event (2016), enabling us to examine foraging habitats under novel and changing climates. First passage time and generalized additive mixed models indicated that during 183 short and 110 long trips (n = 32 birds), wind-assisted flight efficiency, proximity to productive areas, and moonlit-searching were important in both subtropical and subarctic habitats. Laysan albatross took foraging trips that had similar lengths and durations in 2014 and 2016 and visited similar areas, indicating that their foraging range did not expand in response to climatic variability. A strategy that uses similar foraging areas across years combined with reliance on environmental processes that enhance flight efficiency (wind) and that enable searching behaviors (moonlight) indicate that Laysan albatross exhibit complex behavioral plasticity that allows them to utilize subtropical and subarctic habitats affected by dynamic climate variability. This strategy may benefit their ability to respond to oceanographic and climatic change, including expanding warm water regions and changing atmospheric conditions influenced by global warming.

Alaska, Hawaii

Lingering impacts of the 2014-2016 northeast Pacific marine heatwave on seabird demography in Cook Inlet, Alaska (USA)

A protracted period (2014-2016) of anomalously warm water in the northeast Pacific Ocean precipitated an extensive die-off of common murres Uria aalge (hereafter ‘murres’) during 2015-2016, accompanied by reduced colony attendance and reproductive success of murres and black-legged kittiwakes Rissa tridactyla (‘kittiwakes’) starting in 2015. Most murres died of starvation following a large-scale reduction in abundance and quality of forage fish. To assess murre and kittiwake recovery following the marine heatwave, we monitored their demographics at 2 colonies (Chisik and Gull Islands) in Cook Inlet, Alaska (USA), from 2016 to 2019. Compared to historic data (1995-1999), we observed declines and increased variability in colony attendance and productivity across species and colonies, and predation was widespread. At Chisik, where food limitations were common during historic studies, both species experienced substantial population declines and reproductive failures in all 4 years (2016-2019) following the heatwave. At Gull, a typically productive colony during historic studies, murres failed to fledge chicks for 3 years (2016-2018) following the heatwave. By 2019, murre productivity recovered to about half that observed during historic studies (0.28 vs. 0.54 chicks per pair), but populations had declined by half. Kittiwake population size at Gull declined a quarter from historic counts, and reproduction alternated between complete breeding failures (2016/2018) and high productivity (2017/2019). These multi-year demographic impacts indicate lingering effects of the heatwave on kittiwakes and murres through forage fish depletion and increased predator disturbance, and possibly other stressors. It remains unknown whether populations can rebound to historic levels. If so, recovery would likely take decades.

Alaska

Direct and indirect pathways for environmental drivers of hatching success in the loggerhead sea turtle

Nest site selection has consequences for hatching success by mediating the temperature and moisture conditions that eggs experience during the incubation period. Understanding the potentially complex pathways by which nest placement influences these abiotic mediators, and therefore hatching success, is important for predicting which nests will be successful and which may require management action. We studied the effects of loggerhead sea turtle (Caretta caretta) nest site selection on hatching success by linking nest placement characteristics to hatching success through a structural equation model. We monitored 170 nests on Ossabaw Island, Georgia, during the summers of 2017 and 2018 and tracked nest conditions throughout the incubation period. Temperature had a complex effect on hatching success—nests had higher hatching rates if they were exposed to higher mean temperatures but also if they experienced both extremely high (>34°C) and extremely low (<26.5°C) temperatures, suggesting that temperature variability plays a role in determining nest outcomes beyond the mean temperature. Likewise, hatching success declined with a higher incidence of nests being inundated by tides. We found that nests placed at the highest elevations had the highest hatching success rates, likely because those nests had a much lower chance of being washed over by high tides and had higher mean temperatures. Nests were also more successful when placed in greater amounts of vegetation, again because vegetated nests were generally warmer and were associated with fewer washover events. These results shed light on the mechanisms behind selection for certain nest site characteristics and can guide the relocation of nests as a conservation action.

Georgia

Zostera marina (eelgrass) growth and survival along a gradient ofnutrients and turbidity in the lower Chesapeake Bay

Survival of transplanted Zostera marina L. (eelgrass), Z. marina growth,and environmental conditions were studied concurrently at a number of sitesin a southwestern tributary of the Chesapeake Bay to elucidate the factorslimiting macrophyte distribution in this region. Consistent differences insurvival of the transplants were observed, with no long-term survival at anyof the sites that were formerly vegetated with this species but thatcurrently remain unvegetated. Therefore, the current distribution of Z.marina likely represents the extent of suitable environmental conditions inthe region, and the lack of recovery into historically vegetated sites is notsolely due to lack of propagules. Poor long-term survival was related toseasonally high levels of water column light attenuation. Fall transplantsdied by the end of summer following exposure to levels of high springturbidity (K(d) > 3.0). Accumulation of an epiphyte matrix during the latespring (0.36 to 1.14 g g-1 dry wt) may also have contributed to thisstress. Differences in water column nutrient levels among sites during thefall and winter (10 to 15 ??M dissolved inorganic nitrogen and 1 ??Mdissolved inorganic phosphates) had no observable effect on epiphyteaccumulation or macrophyte growth. Salinity effects were minor and there wereno symptoms of disease. Although summertime conditions resulted indepressions in growth, they did not alone limit long-term survival. It issuggested that water quality conditions enhancing adequate seagrass growthduring the spring may be key to long-term Z. marina survival and successfulrecolonization in this region.

Marine Ecology Progress Series

Fecal DNA metabarcoding shows credible short-term prey detections and explains variation in the gut microbiome of two polar bear subpopulations

This study developed and evaluated DNA metabarcoding to identify the presence of pinniped and cetacean prey DNA in fecal samples of East Greenland (EG) and Southern Beaufort Sea (SB) polar bears Ursus maritimus sampled in the spring of 2015-2019. Prey DNA was detected in half (49/92) of all samples, and when detected, ringed seal Pusa hispida was the predominant prey species, identified in 100% (22/22) of EG and 81% (22/27) of SB polar bear samples with prey DNA detected. Bearded seal Erignathus barbatus DNA was found in 19% (5/27) of SB polar bear samples for which prey DNA was detected. Prey DNA detection frequencies and relative abundances were compared to estimates from quantitative fatty acid signature analysis (QFASA) for a subset of SB polar bears. Ringed seal and bearded seal were the main prey identified by both methods, but QFASA also identified 2 cetacean prey species not found by prey DNA. Differences in DNA metabarcoding vs. QFASA results were likely related to the different dietary timescales captured by each approach, i.e. short-term vs. long-term diet, respectively. Prey DNA detection, sex/age class, and subpopulation significantly explained variation in polar bear gut bacterial composition. Polar bear samples with prey DNA detected were associated with higher abundances of the bacterial classes Clostridia and Bacilli and lower abundances of Negativicutes. Fecal DNA metabarcoding is thus useful for identifying recent prey of polar bears, complementing quantitative and likely longer-term QFASA estimates, and may help understand variation in the polar bear gut microbiome.

Marine Ecology Progress Series (MEPS)

Reduction in overwinter body condition and size of Pacific sand lance has implications for piscivorous predators during marine heatwaves

Acute anomalous ocean warming events, including marine heatwaves (MHWs), have significant effects on reproduction and survival of piscivorous seabirds. Additionally, MHWs have negative effects on seabird fish prey, exacerbating these consequences and resulting in population implications for seabirds. We evaluated the relative body condition of Pacific sand lance Ammodytes personatus , an important seabird forage species, in Haro Strait, a highly productive region of southern British Columbia, Canada. We compared body condition and length of fish cohorts that experienced the 2016 MHW year (MHW cohorts) with fish hatched during 3 subsequent post MHW years (2017-2019). Age-0 MHW cohorts had a seasonal decline in body condition in age-0 fish from 100% in the summer to 81% in the winter, while age-1 fish showed a decline from summer-fall highs of 93.5% to wintertime low of 79.5%. In comparison, post MHW cohorts had a winter body condition that was 2-4 times higher than their MHW cohorts. Similar to previous studies in Alaska during the MHW, age-1 fish failed to grow and reach the typical size that distinguishes them from age-0 fish. Poor sand lance condition and growth in winter may explain the ramifications of a warming ocean for top predators, including seabirds and Pacific salmon, which depend on these prey fish in Haro Strait. Our results support the idea that Haro Strait, which is influenced by estuarine circulation resulting in cooler temperatures than surrounding areas, serves as a climate refugium for sand lance populations in summer and provides buffering capacity to ocean climate warming events.

Marine Ecology Progress Series

Assessing individual movement, habitat use, and behavior of non-breeding marine birds in relation to prey availability in the US Atlantic

Resource availability is a key factor driving marine bird movements and distributions, but direct information on prey availability is difficult to obtain at relevant scales. We present novel methods for describing multi-scale trophic associations, combining movement analyses of marine birds with estimates of forage fish surface aggregations from digital aerial survey data and species occupancy from bottom trawl survey data. We analyzed satellite telemetry data from northern gannets Morus bassanus , red-throated loons Gavia stellata , and long-tailed ducks Clangula hyemalis in the US Atlantic during the non-breeding period. Using discrete-time hidden Markov models to distinguish area-restricted (i.e. putative foraging) from transit movements, we examined how environmental factors influence movement, and how forage fish species distributions and surface aggregations influence habitat use by gannets and loons that have greater dietary reliance. Our results suggest that chlorophyll a concentration significantly affected movement behavior across species, highlighting the importance of higher-productivity areas around estuaries during colder months when regional productivity is low. Though variable across species and seasons, spatial cross-correlation analysis revealed that herring species (Family Clupeidae), including Atlantic menhaden Brevoortia tyrannus , may be important resources; it also showed positive spatial correlations with forage fish aggregations. This suggests that prey patch dynamics and factors driving aggregation formation may be as important as species composition. However, spatial patterns were generally low (<0.3), suggesting a mismatch in spatiotemporal resolution, exemplifying the challenges in quantifying trophic relationships in marine systems. Disentangling predator-prey relationships is critical to understanding the mechanisms driving marine bird behavior in rapidly changing marine systems.

Atlantic Ocean

Lactation performance in polar bears is associated with fasting time and energetic state

Females must continually make resource allocation decisions because of fitness trade-offs between self-maintenance and investment in current offspring, yet factors underpinning these decisions are unresolved. Polar bears Ursus maritimus face considerable allocation challenges when seasonal sea-ice melt precludes access to prey for several months, and females rely solely on energy stores to cover their own energetic needs and provision offspring. We tested how female polar bears regulate lactation during onshore fasting (i.e. capital breeding) and determined the consequences of moderated lactation for females and cubs. Overall, milk energy declined, and lactation was more likely to cease with longer time fasting. Lactation was partially mediated by maternal energetic state and depended on litter characteristics. Milk energy declined more sharply with fasting time (~2.6 times more strongly) in females with 2 offspring compared to those with 1. Females with cubs-of-the-year produced higher energy milk than those with yearlings, and their milk energy also increased more strongly with maternal energy density. Milk energy declines benefited females via reduced depletion of maternal energy reserves, but cub growth decreased. Altered lactation investment likely has consequences for both female survival and the fate of offspring, which could scale up to influence population dynamics. Given that Arctic warming means polar bears across much of their range will experience longer periods without access to primary prey, our results underscore how lactation will likely become increasingly compromised.

Marine Ecology Progress Series

Lack of strong responses to the Pacific marine heatwave by benthivorous marine birds indicates importance of trophic drivers

The Pacific marine heatwave (PMH) of 2014-2016 was an intense, long-lasting environmental disturbance expressed throughout the north Pacific. While dramatic consequences of the PMH on pelagic food webs have been well documented, effects on nearshore food webs, i.e., those based on macroalgae primary productivity, benthic invertebrate intermediate consumers, and specialized benthivorous top predators including some marine birds, are not well understood. We conducted summer and winter coastline surveys in two National Parks in the northern Gulf of Alaska from 2006 – 2022. We evaluated changes in abundance of benthivorous marine birds in relation to the heatwave, after accounting for effects of season and region. We also evaluated changes in abundance of nearshore benthic invertebrate prey data to allow specific consideration of a prey-based mechanism for effects of the PMH across food webs. We found that benthivorous marine birds, consisting largely of sea ducks and shorebirds, did not show a strong response to the PMH, unlike significant effects demonstrated by piscivorous birds in pelagic biomes. Unlike extreme reductions in quantity and quality of forage fish documented in other studies, we found that common benthic invertebrate prey abundance remained relatively stable, with only minor increases or decreases, in association with the PMH. Our results support the hypothesis that food availability has a strong mediating effect of the PMH on upper trophic levels across food webs. These findings show how a large-scale environmental perturbation affects biological communities through trophic pathways, provides insight into ecosystem resiliency, and can inform management strategies in the face of persistent climate change.

Alaska

Relating absolute abundance of an estuarine fish to habitat area in an urbanizing environment

Organisms that rely on salt marsh habitat are an important trophic link, helping to maintain estuarine ecosystem productivity. We used GIS to quantify intertidal (assumed salt marsh) area from aerial photographs taken in 1939 and from software-supplied satellite imagery taken in 2021 for tidal creeks in North Carolina (USA) that have experienced minor (<20%), moderate (20-60%), or substantial (>60%) losses of intertidal habitat over the 8 decades. The current (2022) absolute abundance of adult Fundulus heteroclitus , a trophically important resident fish in US Atlantic estuaries, was estimated over each season in each creek by fitting a Lincoln-Petersen model to tag-recapture data. Current abundances of F. heteroclitus were lowest in creeks with the lowest intertidal area. The median and 2.5/97.5 credible intervals of the posterior probability distribution for the slope of a regression model relating current fish abundance to current intertidal area were positive, demonstrating that intertidal area was a meaningful covariate of abundance. Loss of intertidal area in the creeks between 1939 and 2021 ranged from 8 to 93%. The correlation between current intertidal area and historical loss of this habitat was negative and significant (Pearson r = -0.91, p = 0.012). Parameters from the regression relating current abundance to intertidal area were used to estimate historic F. heteroclitus abundances in each creek using GIS-derived estimates of historic intertidal area. Historic abundances were predicted to have been on average (across study creeks) 7.5 times greater in 1939 than in 2022. Reduced abundances, and thus reduced trophic relay by F. heteroclitus to higher-order consumers, can be expected in estuaries that have lost salt marsh due to inter-decadal development.

Marine Ecologly Progressive Series

Metabolic responses to hypoxia of Lycenchelys verrillii (wolf eelpout) and Glyptocephalus cynoglossus (witch flounder): Sedentary bottom fishes of the Hatteras/Virginia middle slope

We collected wolf eelpouts Lycenchelys verrillii and witch flounder Glyptocephalus cynoglossus from Hatteras (North Carolina, USA) and Virginia (USA) Middle Slope sites using a submersible, and made shipboard measurements of their respiration rates and survival in hypoxic (<10% O2 saturation) and anoxic conditions. Both species from the Hatteras site reduced their respiration rates as ambient oxygen decreased, but eelpouts from the Virginia site maintained a constant respiration rate until oxygen saturation dropped below 20%. Moreover, eelpouts from the Hatteras site were significantly more tolerant of hypoxic conditions than fish from the Virginia site and survived anoxia for short periods. These results and our submersible observations of fish behavior support the hypothesis that the Hatteras Middle Slope fauna is exposed to short-term hypoxia events.

Marine Ecology Progress Series