Reproductive biology of the electric lanternfish Electrona risso (Myctophidae) and the bigscale fishes Melamphaes polylepis and Scopelogadus mizolepis (Melamphaidae).

This study was the first to investigate the key reproductive traits of the electric lantern fish Electrona risso (Myctophidae, n = 918) and the bigscale fishes (Melamphaidae) Melamphaes polylepis (n = 260) and Scopelogadus mizolepis (n = 649). Specimens of these mesopelagic species were collected in March and April 2015 in the eastern Central Atlantic (0-24° N, 20-26° W). Sex ratio was not significantly different from 1:1 in E. risso and M. polylepis but significantly skewed toward female dominance in S. mizolepis. Reproductive phases were determined macroscopically and by histological analyses on selected individuals. Female length at 50% maturity (L50 ) was 55.1 mm standard length (LS ) in E. risso, with an observed female maximum length (Lmax ) of 81.2 mm LS . M. polylepis females had an L50 of 40.2 mm LS and an Lmax of 86.7 mm LS . S. mizolepis had an L50 of 46 mm LS and an Lmax of 97.9 mm LS . The three species show histological features of iteroparity, but the E. risso population appears to occur in two year-classes and experience only one spawning season per lifetime in the study region. All three species are batch-spawners. A batch fecundity of 2668 eggs was estimated from one E. risso individual, with a relative batch fecundity of 369 eggs g-1 gonad-free body mass. M. polylepis had a batch fecundity of 1027 eggs and a relative batch fecundity of 149 eggs g-1 (n = 3). S. polylepis had a batch fecundity of 1545 eggs and a relative batch fecundity of 215 eggs g-1 (n = 21). The median gonado-somatic index during the actively spawning phase of E. risso was 4.5, significantly lower than that of M. polylepis (7.5) and S. mizolepis (7.1). No regressing or regenerating phases were observed in this study. Batch-spawning in all three species is suggested to be advantageous to cope with intra-annual variability in food supply and other risks for offspring survival. With what appears to be in effect a (facultative) semelparous strategy in combination with a short life span in E. risso, interannual differences would have a great effect on population dynamics of this species. Knowledge is still lacking on temporal aspects of reproduction such as the duration of the spawning season and the frequency of spawning, as well as age and growth.

Deep demersal fish communities respond rapidly to warming in a frontal region between Arctic and Atlantic waters.

The assessment of climate impact on marine communities dwelling deeper than the well-studied shelf seas has been hampered by the lack of long-term data. For a long time, the prevailing expectation has been that thermal stability in deep ocean layers will delay ecosystem responses to warming. Few observational studies have challenged this view and indicated that deep organisms can respond exceptionally fast to physical change at the sea surface. To address the depth-specific impact of climate change, we investigated spatio-temporal changes in fish community structure along a bathymetry gradient of 150-1500 m between 1998 and 2016 in East Greenland. Here, the Arctic East Greenland Current and the Atlantic Irminger Current meet and mix, representing a sub-Arctic transition zone. We found the strongest signals of community reorganizations at depths between 350 and 1000 m and only weak responses in the shallowest and deepest regions. Changes were in synchrony with atmospheric warming, loss in sea ice and variability in physical sea surface conditions both within our study region and North of the Denmark Strait. These results suggest that interannual variability and long-term climate trends of the larger ecoregion can rapidly affect fish communities down to 1000-m depth through atmospheric ocean coupling and food web interactions.

Length-weight relationships of 55 mesopelagic fishes from the eastern tropical North Atlantic: Across- and within-species variation (body shape, growth stanza, condition factor).

We present estimates of length-weight relationships (LWRs) of 55 mesopelagic fish species of 13 taxonomic families based on data collected in the eastern tropical North Atlantic (ETNA) in March/April 2015. Our data include novel records for 19 species, while for 25 species LWRs are based on the most robust sample sizes, and for 21 species they are based on the most representative size ranges available up to now. In 31 species, body lengths were within the maximum range of body lengths recorded in the area, with new records of maximum lengths for 13 species. Most values for b fell between 2.5 and 3.5 with a mean exponent b of 3.08 (median 3.12) and a mean a of 0.0172 (median 0.0113). Body shape as covariate ('elongated', 'fusiform' and 'short-deep') strongly determined the variation in log a as a function of parameter b. For the mesopelagic fish species investigated, the form factor a3.0 indicated a significant increase of median a3.0 from 'elongated' to 'fusiform' to 'short-deep' body shapes. Large variability existed in parameter b between species of the same taxonomic family. Isometric growth was indicated in only nine species, whereas a positive allometry was suggested in 22 species. Using segmented regression analysis, we investigated ontogenetic variation in LWRs in 30 species. Of these, 20 species showed a breakpoint in LWR, whereby nearly equal numbers exhibited an increase or a decrease in slope following the breakpoint. Seven out of nine species showed significant regional variation in the slope of the relationship of the relative condition factor Krel vs. body length between two or more regions of the ETNA [eastern and western part of the oxygen minimum zone (LO-E, LO-W), northern and central equatorial region (EQ-N, EQ-C)]. A conspicuous pattern was an increase in Krel with body size in the LO-E (in six out of eight species), whereas in the LO-W and the equatorial regions the majority of species showed a related decrease. These findings support the idea that growth patterns in mesopelagic fishes in tropical regions show species-specific ecological niche and life-history adaptations that are finely tuned to small-scale regional environmental conditions. Comparison of our data with those of other studies emphasises that, regarding the small adult sizes of many mesopelagic fish species, estimates of LWR parameters are strongly influenced by sampled size distributions.

Are we ready to track climate-driven shifts in marine species across international boundaries? - A global survey of scientific bottom trawl data.

Marine biota are redistributing at a rapid pace in response to climate change and shifting seascapes. While changes in fish populations and community structure threaten the sustainability of fisheries, our capacity to adapt by tracking and projecting marine species remains a challenge due to data discontinuities in biological observations, lack of data availability, and mismatch between data and real species distributions. To assess the extent of this challenge, we review the global status and accessibility of ongoing scientific bottom trawl surveys. In total, we gathered metadata for 283,925 samples from 95 surveys conducted regularly from 2001 to 2019. We identified that 59% of the metadata collected are not publicly available, highlighting that the availability of data is the most important challenge to assess species redistributions under global climate change. Given that the primary purpose of surveys is to provide independent data to inform stock assessment of commercially important populations, we further highlight that single surveys do not cover the full range of the main commercial demersal fish species. An average of 18 surveys is needed to cover at least 50% of species ranges, demonstrating the importance of combining multiple surveys to evaluate species range shifts. We assess the potential for combining surveys to track transboundary species redistributions and show that differences in sampling schemes and inconsistency in sampling can be overcome with spatio-temporal modeling to follow species density redistributions. In light of our global assessment, we establish a framework for improving the management and conservation of transboundary and migrating marine demersal species. We provide directions to improve data availability and encourage countries to share survey data, to assess species vulnerabilities, and to support management adaptation in a time of climate-driven ocean changes.

Bottom trawl fishing footprints on the world's continental shelves.

Bottom trawlers land around 19 million tons of fish and invertebrates annually, almost one-quarter of wild marine landings. The extent of bottom trawling footprint (seabed area trawled at least once in a specified region and time period) is often contested but poorly described. We quantify footprints using high-resolution satellite vessel monitoring system (VMS) and logbook data on 24 continental shelves and slopes to 1,000-m depth over at least 2 years. Trawling footprint varied markedly among regions: from <10% of seabed area in Australian and New Zealand waters, the Aleutian Islands, East Bering Sea, South Chile, and Gulf of Alaska to >50% in some European seas. Overall, 14% of the 7.8 million-km2 study area was trawled, and 86% was not trawled. Trawling activity was aggregated; the most intensively trawled areas accounting for 90% of activity comprised 77% of footprint on average. Regional swept area ratio (SAR; ratio of total swept area trawled annually to total area of region, a metric of trawling intensity) and footprint area were related, providing an approach to estimate regional trawling footprints when high-resolution spatial data are unavailable. If SAR was ≤0.1, as in 8 of 24 regions, there was >95% probability that >90% of seabed was not trawled. If SAR was 7.9, equal to the highest SAR recorded, there was >95% probability that >70% of seabed was trawled. Footprints were smaller and SAR was ≤0.25 in regions where fishing rates consistently met international sustainability benchmarks for fish stocks, implying collateral environmental benefits from sustainable fishing.

Parasites as biological tags to track an ontogenetic shift in the feeding behaviour of Gadus morhua off West and East Greenland.

Parasites, being an integral part of every ecosystem and trophically transmitted along the food webs, can provide detailed insights into the structure of food webs and can close the information gap between short-term stomach content analyses and long-term fish otolith analyses. They are useful for tracking ontogenetic shifts in the host's diet, the occurrence of specific organisms or migratory behaviour of their hosts, even in inaccessible environments. In the present study, stomach content analyses and parasitological examinations were performed on 70 Atlantic cod Gadus morhua, one of the most important high-level predators of small fish in the North Atlantic, caught during one research vessel cruise from West and East Greenlandic waters. Analyses revealed significant differences in fish size with higher values for East Greenland (average total length (TL) of 50.5 cm) compared to West Greenland (average TL of 33.3 cm). Clear differences were also present in prey and parasite composition. Crustacea was the main food source for all fish (IRI = 10082.70), while the importance of teleosts increased with fish size. With a prevalence of 85 % in West Greenland and 100 % in East Greenland, Nematoda were the most abundant parasite group. The results indicate an ontogenetic shift in the diet, which are discussed in the context of the common distribution theory, stock dynamics and migratory behaviour.

Correction: Community ecology in 3D: Tensor decomposition reveals spatio-temporal dynamics of large ecological communities.

[This corrects the article DOI: 10.1371/journal.pone.0188205.].

Community ecology in 3D: Tensor decomposition reveals spatio-temporal dynamics of large ecological communities.

Understanding spatio-temporal dynamics of biotic communities containing large numbers of species is crucial to guide ecosystem management and conservation efforts. However, traditional approaches usually focus on studying community dynamics either in space or in time, often failing to fully account for interlinked spatio-temporal changes. In this study, we demonstrate and promote the use of tensor decomposition for disentangling spatio-temporal community dynamics in long-term monitoring data. Tensor decomposition builds on traditional multivariate statistics (e.g. Principal Component Analysis) but extends it to multiple dimensions. This extension allows for the synchronized study of multiple ecological variables measured repeatedly in time and space. We applied this comprehensive approach to explore the spatio-temporal dynamics of 65 demersal fish species in the North Sea, a marine ecosystem strongly altered by human activities and climate change. Our case study demonstrates how tensor decomposition can successfully (i) characterize the main spatio-temporal patterns and trends in species abundances, (ii) identify sub-communities of species that share similar spatial distribution and temporal dynamics, and (iii) reveal external drivers of change. Our results revealed a strong spatial structure in fish assemblages persistent over time and linked to differences in depth, primary production and seasonality. Furthermore, we simultaneously characterized important temporal distribution changes related to the low frequency temperature variability inherent in the Atlantic Multidecadal Oscillation. Finally, we identified six major sub-communities composed of species sharing similar spatial distribution patterns and temporal dynamics. Our case study demonstrates the application and benefits of using tensor decomposition for studying complex community data sets usually derived from large-scale monitoring programs.

From Metaphors to Formalism: A Heuristic Approach to Holistic Assessments of Ecosystem Health.

Environmental policies employ metaphoric objectives such as ecosystem health, resilience and sustainable provision of ecosystem services, which influence corresponding sustainability assessments by means of normative settings such as assumptions on system description, indicator selection, aggregation of information and target setting. A heuristic approach is developed for sustainability assessments to avoid ambiguity and applications to the EU Marine Strategy Framework Directive (MSFD) and OSPAR assessments are presented. For MSFD, nineteen different assessment procedures have been proposed, but at present no agreed assessment procedure is available. The heuristic assessment framework is a functional-holistic approach comprising an ex-ante/ex-post assessment framework with specifically defined normative and systemic dimensions (EAEPNS). The outer normative dimension defines the ex-ante/ex-post framework, of which the latter branch delivers one measure of ecosystem health based on indicators and the former allows to account for the multi-dimensional nature of sustainability (social, economic, ecological) in terms of modeling approaches. For MSFD, the ex-ante/ex-post framework replaces the current distinction between assessments based on pressure and state descriptors. The ex-ante and the ex-post branch each comprise an inner normative and a systemic dimension. The inner normative dimension in the ex-post branch considers additive utility models and likelihood functions to standardize variables normalized with Bayesian modeling. Likelihood functions allow precautionary target setting. The ex-post systemic dimension considers a posteriori indicator selection by means of analysis of indicator space to avoid redundant indicator information as opposed to a priori indicator selection in deconstructive-structural approaches. Indicator information is expressed in terms of ecosystem variability by means of multivariate analysis procedures. The application to the OSPAR assessment for the southern North Sea showed, that with the selected 36 indicators 48% of ecosystem variability could be explained. Tools for the ex-ante branch are risk and ecosystem models with the capability to analyze trade-offs, generating model output for each of the pressure chains to allow for a phasing-out of human pressures. The Bayesian measure of ecosystem health is sensitive to trends in environmental features, but robust to ecosystem variability in line with state space models. The combination of the ex-ante and ex-post branch is essential to evaluate ecosystem resilience and to adopt adaptive management. Based on requirements of the heuristic approach, three possible developments of this concept can be envisioned, i.e. a governance driven approach built upon participatory processes, a science driven functional-holistic approach requiring extensive monitoring to analyze complete ecosystem variability, and an approach with emphasis on ex-ante modeling and ex-post assessment of well-studied subsystems.

Terrestrial and Marine Foraging Strategies of an Opportunistic Seabird Species Breeding in the Wadden Sea.

Lesser black-backed gulls Larus fuscus are considered to be mainly pelagic. We assessed the importance of different landscape elements (open sea, tidal flats and inland) by comparing marine and terrestrial foraging behaviours in lesser black-backed gulls breeding along the coast of the southern North Sea. We attached GPS data loggers to eight incubating birds and collected information on diet and habitat use. The loggers recorded data for 10-19 days to allow flight-path reconstruction. Lesser black-backed gulls foraged in both offshore and inland areas, but rarely on tidal flats. Targets and directions were similar among all eight individuals. Foraging trips (n = 108) lasted 0.5-26.4 h (mean 8.7 h), and ranges varied from 3.0-79.9 km (mean 30.9 km). The total distance travelled per foraging trip ranged from 7.5-333.6 km (mean 97.9 km). Trips out to sea were significantly more variable in all parameters than inland trips. Presence in inland areas was closely associated with daylight, whereas trips to sea occurred at day and night, but mostly at night. The most common items in pellets were grass (48%), insects (38%), fish (28%), litter (26%) and earthworms (20%). There was a significant relationship between the carbon and nitrogen isotope signals in blood and the proportional time each individual spent foraging at sea/land. On land, gulls preferentially foraged on bare ground, with significantly higher use of potato fields and significantly less use of grassland. The flight patterns of lesser black-backed gulls at sea overlapped with fishing-vessel distribution, including small beam trawlers fishing for shrimps in coastal waters close to the colony and large beam-trawlers fishing for flatfish at greater distances. Our data show that individuals made intensive use of the anthropogenic landscape and seascape, indicating that lesser black-backed gulls are not a predominantly marine species during the incubation period.

Changes in the seasonal cycles of inorganic nutrients in the coastal zone of the southeastern North Sea from 1960 to 1997: effects of eutrophication and sensitivity to meteoclimatic factors.

A long-term analysis of seasonal cycles of inorganic nutrients by means of a seasonal index is presented for the German Bight and the southern Wadden Sea (SE North Sea). Multivariate analysis for the German Bight data series revealed dependence of ammonium and phosphate index time series on dissolved inorganic nitrogen concentrations and riverine nutrient loads. Both indices are assumed to reflect seasonal dynamics of remineralisation processes associated with increased supply of organic matter. Temporal analysis revealed breaks in nutrient dynamics in 1970/1972 and 1979/1980. After 1970/1972, an unprecedented increase in the summer concentrations of mineralisation endproducts with correspondingly low index values were observed, which further declined after 1979/1980. Further breaks for Wadden Sea data series were identified in 1985 and 1988/1989. The indicative value of the temporal breaks with respect to eutrophication is discussed against the background of changes in meteoclimatic factors and local environmental conditions. Collated information on eutrophication effects was consistent with the observed breaks. It is suggested that the break in 1970/1972 in the German Bight was the first sign of ecosystem response to eutrophication in the SE North Sea.