Deep Pelagic Waters Research Articles

Ecosystem Metabolism and Gradients of Temperature, Oxygen and Dissolved Inorganic Carbon in the Littoral Zone of a Macrophyte‐Dominated Lake

AbstractDense submerged macrophyte stands in lakes may promote alternating daytime stratification and nighttime convective mixing, driving extensive spatiotemporal variations in water temperature, oxygen, pH, and inorganic carbon (DIC). We set out to investigate environmental conditions and ecosystem metabolism in the macrophyte‐dominated littoral zone (0.6 m depth) of a shallow, mesotrophic lake and compared this with the pelagic zone (3.0 m deep). We found that, during summer, vertical water column gradients only occasionally occurred and were weak in the pelagic zone, while dense littoral macrophyte stands of charophytes exhibited strong diel changes and steep daytime temperature and oxygen depth gradients. Oxygen showed daytime surface supersaturation and bottom anoxia, alternating with nighttime mixing. In spring, before charophytes appeared, the vertical gradients were largely absent. Ecosystem metabolism was primarily positive in spring, but areal rates of daily gross primary production (GPP) and closely related respiration increased 6‐fold in summer. Ecosystem metabolism calculated based on oxygen or DIC was very similar, with a 1:1 M basis. Daytime DIC loss by CaCO3precipitation on charophyte surfaces averaged 6.3% of total DIC loss, and this loss was restored during nighttime. The contribution of shallow littoral macrophyte communities to whole‐lake summer GPP exceeded that estimated for phytoplankton. Environmental conditions in the macrophyte‐covered littoral zone differed markedly from pelagic waters with steep vertical gradients in temperature and chemistry during daytime including anoxic micro‐habitats in stark contrast to deep pelagic waters. The study highlights the significant influence of the littoral zone and macrophytes on whole‐lake ecosystem processes.

ROV observations reveal infection dynamics of gill parasites in midwater cephalopods

Gill parasites of coleoid cephalopods are frequently observed during remotely operated vehicle (ROV) dives in the Monterey Submarine Canyon. However, little knowledge exists on the identity of the parasite species or their effects on the cephalopod community. With the help of ROV-collected specimens and in situ footage from the past 27 years, we report on their identity, prevalence and potential infection strategy. Gill parasites were genetically and morphologically identified from collected specimens of Chiroteuthis calyx, Vampyroteuthis infernalis and Gonatus spp. In situ prevalence was estimated from video footage for C. calyx, Galiteuthis spp., Taonius spp. and Japetella diaphana, enabled by their transparent mantle tissue. The most common parasite was identified as Hochbergia cf. moroteuthensis, a protist of unresolved taxonomic ranking. We provide the first molecular data for this parasite and show a sister group relationship to the dinoflagellate genus Oodinium. Hochbergia cf. moroteuthensis was most commonly observed in adult individuals of all species and was sighted year round over the analyzed time period. In situ prevalence was highest in C. calyx (75%), followed by Galiteuthis spp. (29%), Taonius spp. (27%) and J. diaphana (7%). A second parasite, not seen on the in situ footage, but occurring within the gills of Gonatus berryi and Vampyroteuthis infernalis, could not be found in the literature or be identified through DNA barcoding. The need for further investigation is highlighted, making this study a starting point for unravelling ecological implications of the cephalopod-gill-parasite system in deep pelagic waters.

Seasonality of density currents induced by differential cooling

Abstract. When lakes experience surface cooling, the shallow littoral region cools faster than the deep pelagic waters. The lateral density gradient resulting from this differential cooling can trigger a cold downslope density current that intrudes at the base of the mixed layer during stratified conditions. This process is known as a thermal siphon (TS). TSs flush the littoral region and increase water exchange between nearshore and pelagic zones; thus, they may potentially impact the lake ecosystem. Past observations of TSs in lakes are limited to specific cooling events. Here, we focus on the seasonality of TS-induced lateral transport and investigate how seasonally varying forcing conditions control the occurrence and intensity of TSs. This research interprets 1-year-long TS observations from Rotsee (Switzerland), a small wind-sheltered temperate lake with an elongated shallow region. We demonstrate that TSs occur for more than 50 % of the days from late summer to winter and efficiently flush the littoral region within ∼10 h. We further quantify the occurrence, intensity, and timing of TSs over seasonal timescales. The conditions for TS formation become optimal in autumn when the duration of the cooling phase is longer than the time necessary to initiate a TS. The decrease in surface cooling by 1 order of magnitude from summer to winter reduces the lateral transport by a factor of 2. We interpret this transport seasonality with scaling relationships relating the daily averaged cross-shore velocity, unit-width discharge, and flushing timescale to the surface buoyancy flux, mixed-layer depth, and lake bathymetry. The timing and duration of diurnal flushing by TSs relate to daily heating and cooling phases. The longer cooling phase in autumn increases the flushing duration and delays the time of maximal flushing relative to the summer diurnal cycle. Given their scalability, the results reported here can be used to assess the relevance of TSs in other lakes and reservoirs.

The vertical distribution and biological transport of marine microplastics across the epipelagic and mesopelagic water column

Plastic waste has been documented in nearly all types of marine environments and has been found in species spanning all levels of marine food webs. Within these marine environments, deep pelagic waters encompass the largest ecosystems on Earth. We lack a comprehensive understanding of the concentrations, cycling, and fate of plastic waste in sub-surface waters, constraining our ability to implement effective, large-scale policy and conservation strategies. We used remotely operated vehicles and engineered purpose-built samplers to collect and examine the distribution of microplastics in the Monterey Bay pelagic ecosystem at water column depths ranging from 5 to 1000 m. Laser Raman spectroscopy was used to identify microplastic particles collected from throughout the deep pelagic water column, with the highest concentrations present at depths between 200 and 600 m. Examination of two abundant particle feeders in this ecosystem, pelagic red crabs (Pleuroncodes planipes) and giant larvaceans (Bathochordaeus stygius), showed that microplastic particles readily flow from the environment into coupled water column and seafloor food webs. Our findings suggest that one of the largest and currently underappreciated reservoirs of marine microplastics may be contained within the water column and animal communities of the deep sea.

Far away from home: presence of fur seal (Arctocephalus sp.) in the equatorial Atlantic Ocean

The occurrence of pinnipeds in areas distant from their reproductive sites is a well-known phenomenon, which could be influenced by oceanographic anomalies in a changing environment. In the present paper, we present an extra-limital record in deep pelagic waters of a young fur seal (Arctocephalus sp.) in the North Atlantic Ocean. The individual was recorded alive in March/April 2013 in the vicinity of the Sao Pedro and Sao Paulo Archipelago (ca. 00°56′N; 29°22′W), which is about 1010 km away from the Brazilian mainland, just above the equatorial line. The specimen was apparently healthy, and it was feeding upon flying fish (Cheilopogon cyanopterus). We hypothesized it to be one of the three main potential species (Arctocephalus tropicalis, Arctocephalus australis, and Arctocephalus pusillus pusillus), suggested their possible routes of arrival, and highlighted it as the northernmost record of a fur seal in the Atlantic Ocean.

Relationship between cetacean presence, their prey, and oceanographic conditions at an offshore pelagic environment

Large mobile predators take advantage of offshore pelagic environments that tend to be surveyed at a low temporal scale. We present results from a one-year observation of passive and active acoustics, as well as oceanographic measures, taken at a moored location 150 nmi offshore southern California in 4000 m deep pelagic waters. The passive acoustic data shows presence of delphinids and baleen, sperm, and beaked whales throughout the recording period with species-specific seasonal occurrence. Variability in small pelagics, krill, and diel vertical migrators is evident from the active acoustic data. Multiple upwelling events and associated changes in dissolved oxygen at 500 m water depth, as well as in surface waters, were also recorded over the year. We will present results on the relationships among oceanographic variability, presence of pelagic species, and top predator presence. This study will provide valuable information on ecosystem processes in meso- and epipelagic waters on a relatively small spatial scale but very fine temporal scale.

Feasibility of using the National Marine Mammal Tissue Bank for retrospective exploratory studies of perfluorinated alkyl acids

Perfluorinated alkyl acids (PFAAs) have been used for 50+ years in materials such as stain-resistant treatments for paper and clothing, lubricants, and foam fire extinguishers. PFAAs are characterized by a fully fluorinated alkyl chain with a terminal acid group. Their long half-lives and ubiquitous environmental distribution create considerable concern for wildlife and human exposure. There is interest in examining temporal trends of PFAAs using the National Marine Mammal Tissue Bank (NMMTB), but NMMTB tissues are frozen and cryohomogenized in polytetrafluoroethylene (PTFE)-based materials. Because PTFE supplies may leach PFAAs into samples, this study mimicked collection, processing and storage steps of NMMTB samples and measured PFAA leaching to determine the feasibility of using this sample archive for PFAA temporal trends. We also explored concentrations in Atlantic white-sided dolphin (Lagenorhynchus acutus, WSDs) and rough-toothed dolphin (Steno bredanensis, RTDs) blubber (n=3 and 0) and liver (n=48 and 12, respectively). The materials used in NMMTB protocols may add up to 0.968ng/g perfluorooctanoic acid (PFOA), 0.090ng/g perfluorononanoic acid (PNFA), and 0.221ng/g perfluorooctane sulfonate (PFOS) to each archived sample. Leaching of PFNA and PFOS from supplies compared to dolphin levels was negligible, but PFOA contributions were substantially higher than levels found in most dolphin liver samples. Therefore, monitoring PFOA temporal trends from the NMMTB would require careful consideration. RTDs had significantly higher levels of PFOS and PFNA than WSDs. Both species have similar life history, trophic status, and foraging behaviors in deep pelagic waters, so differences could be from latitudinal variation in contamination. RTDs stranded in Florida; WSDs stranded farther north mostly in Massachusetts. Juveniles had significantly higher levels of PFOS and PFNA than adults in both species, suggesting growth dilution as they approach maturity. PFOS significantly decreased after 2001 in both species as expected based on changes in production.

Behavioral responses to annual temperature variation alter the dominant energy pathway, growth, and condition of a cold-water predator

There is a pressing need to understand how ecosystems will respond to climate change. To date, no long-term empirical studies have confirmed that fish populations exhibit adaptive foraging behavior in response to temperature variation and the potential implications this has on fitness. Here, we use an unparalleled 11-y acoustic telemetry, stable isotope, and mark-recapture dataset to test if a population of lake trout (Salvelinus namaycush), a cold-water stenotherm, adjusted its use of habitat and energy sources in response to annual variations in lake temperatures during the open-water season and how these changes translated to the growth and condition of individual fish. We found that climate influenced access to littoral regions in spring (data from telemetry), which in turn influenced energy acquisition (data from isotopes), and growth (mark-recapture data). In more stressful years, those with shorter springs and longer summers, lake trout had reduced access to littoral habitat and assimilated less littoral energy, resulting in reduced growth and condition. Annual variation in prey abundance influenced lake trout foraging tactics (i.e., the balance of the number and duration of forays) but not the overall time spent in littoral regions. Lake trout greatly reduced their use of littoral habitat and occupied deep pelagic waters during the summer. Together, our results provide clear evidence that climate-mediated behavior can influence the dominant energy pathways of top predators, with implications ranging from individual fitness to food web stability.

Long-Term Monitoring of Dolphin Biosonar Activity in Deep Pelagic Waters of the Mediterranean Sea

Dolphins emit short ultrasonic pulses (clicks) to acquire information about the surrounding environment, prey and habitat features. We investigated Delphinidae activity over multiple temporal scales through the detection of their echolocation clicks, using long-term Passive Acoustic Monitoring (PAM). The Istituto Nazionale di Fisica Nucleare operates multidisciplinary seafloor observatories in a deep area of the Central Mediterranean Sea. The Ocean noise Detection Experiment collected data offshore the Gulf of Catania from January 2005 to November 2006, allowing the study of temporal patterns of dolphin activity in this deep pelagic zone for the first time. Nearly 5,500 five-minute recordings acquired over two years were examined using spectrogram analysis and through development and testing of an automatic detection algorithm. Echolocation activity of dolphins was mostly confined to nighttime and crepuscular hours, in contrast with communicative signals (whistles). Seasonal variation, with a peak number of clicks in August, was also evident, but no effect of lunar cycle was observed. Temporal trends in echolocation corresponded to environmental and trophic variability known in the deep pelagic waters of the Ionian Sea. Long-term PAM and the continued development of automatic analysis techniques are essential to advancing the study of pelagic marine mammal distribution and behaviour patterns.

Food web properties of the recently constructed, deep subtropical Fei-Tsui Reservoir in comparison with the ancient Lake Biwa

Using carbon and nitrogen stable isotope analysis, we characterised food web properties of the deep subtropical Fei-Tsui Reservoir (FTR), which was recently altered from a lotic to a lentic system after dam construction. In the littoral zone, zoobenthos showed strong reliance (83.9%) on benthic algal production. Zoobenthos were never found in the profundal zone because of anoxia. Zooplankton depleted 13C more than that of particulate organic matter as their putative food source, suggesting a contribution of methane-derived carbon to pelagic food webs. Excluding juveniles, non-native and fluvial species, adult fish showed strong reliance (on average 80.9%) on benthic production, weakly coupled with pelagic food webs. These results contrast low benthic production reliance (on average 27.4%) for a fish community in Lake Biwa, which is also classified as a subtropical lake. Both lakes are characterised by deep pelagic waters but quite different in their geological ages, suggesting that the aquatic communities in the FTR have fluvial origins, and their lacustrine history was too short for them to adapt to newly emerged deep pelagic habitat. Our isotope data are useful as a reference of newly established lentic food webs to monitor ongoing ecological and evolutionary dynamics as a result of anthropogenic disturbances.

Long-term monitoring of cetacean bioacoustics using cabled observatories in deep-sea off East Sicily

The EMSO Research Infrastructure operates multidisciplinary seafloor-cabled observatories in a deep-sea area offshore Eastern Sicily (2100 m of depth). In a data-lacking zone, Passive Acoustic Monitoring activities revealed new information on cetacean bioacoustics over multiple ecological scales. Expert operators investigated the presence of cetacean vocalizations within the large acoustic datasets acquired. Then, algorithms were developed to provide information on the behavior and ecology of the recorded species. In 2005-2006, the acoustic activity of toothed whales was investigated through the OvDE antenna (100 Hz to 48 kHz). The assessment of the size distribution of sperm whales was acoustically possible and the tracking of the animals showed the direction of movement and the diving profile. The biosonar activity of dolphins resulted mostly confined in the nighttime, linked to seasonal variation in daylight time and prey-field variability known for these deep-pelagic waters. Furthermore, in 2012-2013, we monitored the annual acoustic presence of fin whales thanks to the NEMO-SN1 station (1 Hz to 1 kHz). The results showed that the species was present throughout all seasons, with peaks in call detection rate during spring and summer months, and that the fin whale calls were mostly detected in low background noise conditions.

Interpopulation resource partitioning of Lesser Frigatebirds and the influence of environmental context

Conspecific individuals inhabiting nearby breeding colonies are expected to compete strongly for food resources owing to the constraints imposed by shared morphology, physiology, and behavior on foraging strategy. Consequently, colony‐specific foraging patterns that effectively partition the available resources may be displayed. This study aimed to determine whether intraspecific resource partitioning occurs in two nearby colonies of Lesser Frigatebirds (Fregata ariel). A combination of stable isotope analysis and GPS tracking was used to assess dietary and spatial partitioning of foraging resources during the 2013 and 2014 breeding seasons. These results were compared to vessel‐derived estimates of prey availability, local primary productivity, and estimates of reproductive output to suggest potential drivers and implications of any observed partitioning. Isotopic data indicated a more neritic source of provisioned resources for near‐fledged chicks at an inshore colony, whereas their offshore counterparts were provisioned with resources with a more pelagic signal. Deep pelagic waters (>200 m) had higher availability of a preferred prey type despite a trend for lower primary productivity. Differences in foraging ecology between the two populations may have contributed to markedly different reproductive outputs. These findings suggest environmental context influences dietary and spatial aspects of foraging ecology. Furthermore, the effect of colony‐specific foraging patterns on population demography warrants further research.

Habitat use of hourglass dolphins near the South Shetland Islands, Antarctica

The hourglass dolphin Lagenorhynchus cruciger is the only regularly occurring small delphinid found south of the Antarctic Polar Front, yet little is known about its ecology and habitat use. This study uses 8 years (14 cruises) of standardized shipboard surveys during January–March (2003–2011) in southern Drake Passage near the South Shetland Islands to summarize the spatial distribution of hourglass dolphin sightings and quantify habitat use. Sighting data are linked to bathymetry (depth, slope) and distance to the average location of oceanographic features. A generalized linear model is used to examine the relationships between sightings and habitat features. Hourglass dolphins were sighted on 50% of surveys (n = 29); sightings were concentrated in February. Group size tended to be 2–6 individuals; there were only 2 sightings of larger groups, of 15 and 25 individuals. Sightings were distributed entirely within the deep pelagic waters north of the South Shetland Islands in southern Drake Passage and were closely associated with the southern boundary of the Antarctic Circumpolar Current. Information on occurrence and distribution reported in this study may be useful for refining habitat associations for hourglass dolphins at regional scales in the Southern Ocean.

Encounters with pelagic and continental slope cetacean species near the northern shore of the Gulf of Taranto, Italy

Little is known about the distribution and species composition of cetaceans in the Gulf of Taranto, central Mediterranean Sea. Available information is limited to stranding reports and to scant preliminary observations at sea. Cetacean sightings were recorded between April 2007 and October 2009 during 23 daily boat surveys, totalling 1728 km of navigation. Three cetacean species were encountered: striped dolphins (n = 41), Risso's dolphins (n = 5) and fin whales (n = 3). An additional sighting of a sperm whale group is reported. Encounter rates of striped dolphins were the highest, suggesting a regular occurrence in the study area. Risso's dolphins seemed to be predictably present in continental slope waters off the coast of Basilicata. Fin whales and striped dolphins, normally inhabiting deep pelagic waters offshore, were found in continental slope waters relatively close to the coast, possibly due to the particular bathymetry of the Gulf of Taranto and to oceanographic conditions that enhance local productivity. High human impact and worsening environmental conditions in the northern Gulf of Taranto raise concerns over the future of cetaceans living in these waters. This study expands previous knowledge and aims to stimulate research on whales and dolphins in what seems to be an important portion of their Mediterranean range.