Distribution of wild and stocked Japanese eels in the lower reaches of the Tone River catchment revealed by otolith stable-isotope ratios.

The natural occurrence of Japanese eels Anguilla japonica in Japan's highly altered catchments is a critical measure of habitat restoration and ecosystem health, yet this metric may be obscured by the incidence of co-occurring cultured eels. Distribution of wild and stocked A. japonica in the lower reaches of the Tone River cachment was investigated using otolith oxygen and carbon stable-isotope ratios (δ18 O and δ13 C). We developed a discrimination model to classify wild and cultured eels for 560 individuals and applied it to eels captured in the lower reaches of the Tone River catchment. Wild eels were found at all study sites of this part of the catchment, indicating natural recruitment of the species occurred from the estuary to third-order streams at least as far as Lake Imbanuma. The proportion of wild eels ranged from 86 to 100% at river and stream sites and was only 23% at upstream Lake Imbanuma sites. This difference probably depends on whether or not eels were released near the corresponding site. More than 50% of stocked eels occurred at sites proximate to release locations, indicating limited dispersal by cultured eels.

Integrating machine learning with otolith isoscapes: Reconstructing connectivity of a marine fish over four decades.

Stable isotopes are an important tool to uncover animal migration. Geographic natal assignments often require categorizing the spatial domain through a nominal approach, which can introduce bias given the continuous nature of these tracers. Stable isotopes predicted over a spatial gradient (i.e., isoscapes) allow a probabilistic and continuous assignment of origin across space, although applications to marine organisms remain limited. We present a new framework that integrates nominal and continuous assignment approaches by (1) developing a machine-learning multi-model ensemble classifier using Bayesian model averaging (nominal); and (2) integrating nominal predictions with continuous isoscapes to estimate the probability of origin across the spatial domain (continuous). We applied this integrated framework to predict the geographic origin of the Northwest Atlantic mackerel (Scomber scombrus), a migratory pelagic fish comprised of northern and southern components that have distinct spawning sites off Canada (northern contingent) and the US (southern contingent), and seasonally overlap in the US fished regions. The nominal approach based on otolith carbon and oxygen stable isotopes (δ13C/δ18O) yielded high contingent classification accuracy (84.9%). Contingent assignment of unknown-origin samples revealed prevalent, yet highly varied contingent mixing levels (12.5-83.7%) within the US waters over four decades (1975-2019). Nominal predictions were integrated into mackerel-specific otolith oxygen isoscapes developed independently for Canadian and US waters. The combined approach identified geographic nursery hotspots in known spawning sites, but also detected geographic shifts over multi-decadal time scales. This framework can be applied to other marine species to understand migration and connectivity at a high spatial resolution, relevant to management of unit stocks in fisheries and other conservation assessments.

Multi-decadal trends in contingent mixing of Atlantic mackerel (Scomber scombrus) in the Northwest Atlantic from otolith stable isotopes.

The Atlantic mackerel (Scomber scombrus) in the Northwest Atlantic is comprised of northern and southern components that have distinct spawning sites off Canada (northern contingent) and the US (southern contingent), and seasonally overlap in US fished regions. Thus, assessment and management of this population can be sensitive to levels of mixing between contingents, which remain unknown. Multi-decadal trends in contingent mixing levels within the US fisheries region were assessed, and the contingent composition across seasons, locations, ages, and size classes were characterized using archived otoliths and developing a classification baseline based on juvenile otolith carbon and oxygen stable isotopes (δ13C/δ18O values). Classification of age ≥ 2 adults demonstrated that northern contingent mixing was prevalent within the US continental shelf waters during the past 2 decades (2000-2019), providing an important seasonal subsidy to the US winter fishery despite substantial depletion in spawning stock biomass of the dominant northern contingent. While the majority of older fish were of the northern contingent during the early 2000s, the southern contingent contribution increased with age/size class during the recent period (2013-2019). Spatial mixing was most prevalent during February and March when the northern contingent occurred as far south as the Delmarva Peninsula, but were mostly absent from US waters in May. A positive relationship (albeit not significant; r = 0.60, p = 0.07) occurred between northern contingent mixing and US fisheries landings, which could imply that higher contingent mixing levels might be associated with greater landings for the US winter mackerel fishery. The yield of the Northwest Atlantic mackerel depends upon the status of the northern contingent, with the southern contingent possibly more prone to depletion. Spatially explicit stock assessment models are recommended to conserve both productivity and stability in this two-component population.

The recurring impact of storm disturbance on black sea bass (Centropristis striata) movement behaviors in the Mid-Atlantic Bight.

Storm events are a significant source of disturbance in the Middle Atlantic Bight, in the Northwest Atlantic, that cause rapid destratification of the water column during the late summer and early fall. Storm-driven mixing can be considered as a seasonal disturbance regime to demersal communities, characterized by the recurrence of large changes in bottom water temperatures. Black sea bass are a model ubiquitous demersal species in the Middle Atlantic Bight, as their predominantly sedentary behavior makes them ideal for tagging studies while also regularly exposing them to summer storm disturbances and the physiological stresses associated with thermal destratification. To better understand the responsiveness of black sea bass to storm impacts, we coupled biotelemetry with a high-resolution Finite Volume Community Ocean Model (FVCOM). During the summers of 2016-2018, 8-15 black sea bass were released each year with acoustic transponders at three reef sites, which were surrounded by data-logging receivers. Data were analyzed for activity levels and reef departures of black sea bass, and fluctuations in temperature, current velocity, and turbulent kinetic energy. Movement rates were depressed with each consecutive passing storm, and late-season storms were associated with permanent evacuations by a subset of tagged fish. Serial increases in bottom temperature associated with repeated storm events were identified as the primary depressor of local movement. Storm-driven increases in turbulent kinetic energy and current velocity had comparatively smaller, albeit significant, effects. Black sea bass represents both an important fishery resource and an indicator species for the impact of offshore wind development in the United States. Their availability to fisheries surveys and sensitivity to wind turbine impacts will be biased during periods of high storm activity, which is likely to increase with regional climate change.

Multiple spawning run contingents and population consequences in migratory striped bass Morone saxatilis.

Multiple spawning run contingents within the same population can experience varying demographic fates that stabilize populations through the portfolio effect. Multiple spawning run contingents (aka run timing groups) are reported here for the first time for striped bass, an economically important coastal species, which is well known for plastic estuarine and shelf migration behaviors. Adult Hudson River Estuary striped bass (n = 66) were tagged and tracked with acoustic transmitters from two known spawning reaches separated by 90 km. Biotelemetry recaptures for two years demonstrated that each river reach was associated with separate contingents. Time series of individual spawning phenologies were examined via nonparametric dynamic time warping and revealed two dominant time series centroids, each associated with a separate spawning reach. The lower spawning reach contingent occurred earlier than the higher reach contingent in 2017 but not in 2018. The majority (89%) of returning adults in 2018 showed the same contingent behaviors exhibited in 2017. Spawning contingents may have been cued differently by temperatures, where warming lagged 1-week at the higher reach in comparison to the lower reach. The two contingents exhibited similar Atlantic shelf migration patterns with strong summer fidelity to Massachusetts Bay and winter migrations to the southern US Mid-Atlantic Bight. Still, in 2017, differing times of departure into nearby shelf waters likely caused the early lower reach contingent to experience substantially higher mortality than the later upper reach contingent. Anecdotal evidence suggests that higher fishing effort is exerted on the early-departing individuals as they first enter shelf fisheries. Thus, as in salmon, multiple spawning units can lead to differential demographic outcomes, potentially stabilizing overall population dynamics.

Differential migration in Chesapeake Bay striped bass.

Differential migration-increased migration propensity with increasing individual size-is common in migratory species. Like other forms of partial migration, it provides spatial buffering against regional differences in habitat quality and sources of mortality. We investigated differential migration and its consequences to survival and reproductive patterns in striped bass, a species with well-known plasticity in migration behaviors. A size-stratified sample of Potomac River (Chesapeake Bay) Morone saxatilis striped bass was implanted with acoustic transmitters and their subsequent coastal shelf migrations recorded over a 4-yr period by telemetry receivers throughout the Mid-Atlantic Bight and Southern New England. A generalized linear mixed model predicted that ≥ 50% of both males and females depart the Chesapeake Bay at large sizes >80 cm total length. Egressing striped bass exited through both the Chesapeake Bay mouth and Delaware Bay (via the Chesapeake and Delaware Canal), favoring the former. All large fish migrated to Massachusetts shelf waters and in subsequent years repeatedly returned to regions within Massachusetts and Cape Cod Bays. Within this dominant behavior, minority behaviors included straying, skipped spawning, and residency by large individuals (those expected to migrate). Analysis of the last day of transmission indicated that small resident striped bass experienced nearly 2-fold higher loss rates (70% yr-1) than coastal shelf emigrants (37% yr-1). The study confirmed expectations for a threshold size at emigration and different mortality levels between Chesapeake Bay (resident) and ocean (migratory) population contingents; and supported the central premise of the current assessment and management framework of a two-contingent population: smaller Chesapeake Bay residents and a larger ocean contingent. An improved understanding of differential migration thus affords an opportunity to specify stock assessments according to different population sub-components, and tailor reference points and control rules between regions and fishing stakeholder groups.

Comparative migration ecology of striped bass and Atlantic sturgeon in the US Southern mid-Atlantic bight flyway.

Seasonal migrations are key to the production and persistence of marine fish populations but movements within shelf migration corridors or, "flyways", are poorly known. Atlantic sturgeon and striped bass, two critical anadromous species, are known for their extensive migrations along the US Mid-Atlantic Bight. Seasonal patterns of habitat selection have been described within spawning rivers, estuaries,and shelf foraging habitats, but information on the location and timing of key coastal migrations is limited. Using a gradient-based array of acoustic telemetry receivers, we compared the seasonal incidence and movement behavior of these species in the near-shelf region of Maryland, USA. Atlantic sturgeon incidence was highest in the spring and fall and tended to be biased toward shallow regions, while striped bass had increased presence during spring and winter months and selected deeper waters. Incidence was transient (mean = ~2 d) for both species with a pattern of increased residency (>2 d) during autumn and winter, particularly for striped bass, with many individuals exhibiting prolonged presence on the outer shelf during winter. Flyways also differed spatially between northern and southern migrations for both species and were related to temperature: striped bass were more likely to occur in cool conditions while Atlantic sturgeon preferred warmer temperatures. Observed timing and spatial distribution within the Mid-Atlantic flyway were dynamic between years and sensitive to climate variables. As shelf ecosystems come under increasing maritime development, gridded telemetry designs represent a feasible approach to provide impact responses within key marine flyways like those that occur within the US Mid-Atlantic Bight.

Population connectivity of pelagic megafauna in the Cuba-Mexico-United States triangle.

The timing and extent of international crossings by billfishes, tunas, and sharks in the Cuba-Mexico-United States (U.S.) triangle was investigated using electronic tagging data from eight species that resulted in >22,000 tracking days. Transnational movements of these highly mobile marine predators were pronounced with varying levels of bi- or tri-national population connectivity displayed by each species. Billfishes and tunas moved throughout the Gulf of Mexico and all species investigated (blue marlin, white marlin, Atlantic bluefin tuna, yellowfin tuna) frequently crossed international boundaries and entered the territorial waters of Cuba and/or Mexico. Certain sharks (tiger shark, scalloped hammerhead) displayed prolonged periods of residency in U.S. waters with more limited displacements, while whale sharks and to a lesser degree shortfin mako moved through multiple jurisdictions. The spatial extent of associated movements was generally associated with their differential use of coastal and open ocean pelagic ecosystems. Species with the majority of daily positions in oceanic waters off the continental shelf showed the greatest tendency for transnational movements and typically traveled farther from initial tagging locations. Several species converged on a common seasonal movement pattern between territorial waters of the U.S. (summer) and Mexico (winter).

High resolution micromill sampling for analysis of fish otoliths by ICP-MS: effects of sampling and specimen preparation on trace element fingerprints.

Otoliths are calcified structures in the head of fish that record environmental information about fish's life history. Gathering the elemental information from the core of an otolith corresponding to the juvenile period of fish's life is critical to discriminate the adult fish to their natal habitats reliably. A high resolution micromill has been used to isolate the otolith core from a whole otolith for elemental analysis. The effects of micromilling procedures (e.g., sectioning, embedding and drilling) on contamination to otolith trace element levels were examined using paired blackfin tuna (Thunnus atlanticus) otoliths. Otoliths were decontaminated by dilute hydrogen peroxide and nitric acid throughout to remove surface contamination. A preconcentration procedure was used to determine the trace elements from the small core material by ICP-MS. It was found that micromilling procedures introduce significant contamination to otoliths, especially for Al, Cu, Pb and Zn. The sectioning procedure caused significant contamination for Co and Cu, while the embedding procedure resulted in contamination for nearly all trace elements (Al, Cd, Co, Cu, Ga, Mn, Ni, Pb, V and Zn). The combined sectioning, embedding and drilling procedure also resulted in contamination for most trace elements. Despite the contamination across all procedural steps, the decontamination procedure effectively removed the surface contamination with the exception of Pb and Zn. Bias (e.g., residual contamination) on Pb was small in comparison to overall concentration of Pb expected to occur in fish otoliths, therefore, its effect may be minor in discriminating individuals. Bias on Zn was larger that could limit its application in discriminating individuals.

Coastal evacuations by fish during extreme weather events.

An increase in the intensity and frequency of extreme events is predicted to occur as a result of climate change. In coastal ecosystems, hurricanes and flooding can cause dramatic changes in water quality resulting in large mortality events in estuarine fauna. Facultative migration behaviors represent a key adaptation by which animals can evacuate ecological catastrophes, but remain poorly studied in marine systems. Here we identify coastal evacuations by otherwise resident riverine striped bass in the Hudson River Estuary, New York, USA, caused by an intense period of tropical storms in autumn 2011. These storms produced record rainfall and high water discharges into the Hudson River Estuary that increased the water level and reduced the water temperature, salinity and dissolved oxygen levels. Striped bass moved out of the estuary, exhibiting novel migration behaviours, that may have been in response to the strong flow and unsuitable conditions. In the months following the storms, some fish demonstrated exploratory trips back to the estuary, which may have been to assess the conditions before returning for the remainder of the winter. Behavioural adaptions to weather events by striped bass and other coastal fishes will depend on maintenance of key population segments and unimpeded evacuation routes.

Atlantic bluefin tuna (Thunnus thynnus) population dynamics delineated by organochlorine tracers.

Atlantic bluefin tuna (ABFT) are highly valued and heavily exploited, and critical uncertainties regarding their population structure hinder effective management. Evidence supports the existence of two breeding populations of ABFT; a western population in the Gulf of Mexico and an eastern population in the Mediterranean Sea; both of which migrate and mix in the North Atlantic. Conventional tagging studies suggest low rates of trans-Atlantic migrations; however, electronic tagging and stable isotopes in otoliths indicate stock mixing up to 57% between management zones delineated by 45 degrees W longitude. Here we show that organochlorine pesticides and polychlorinated biphenyls (PCBs) can be used as tracers of bluefin tuna foraging grounds in the North Atlantic and confirm that stock mixing of juvenile tuna within the U.S. Mid Atlantic Bight is indeed high (33-83% eastern origin), and is likely spatially and temporally variable. We further demonstrate that >10% of the Mediterranean population is migratory, that young bluefin tuna migrate from the Mediterranean to western Atlantic foraging grounds as early as age 1, and then return to the Mediterranean Sea as young as age 5, presumably to breed. The tracer method described here provides a novel means for distinguishing bluefin tuna populations and ontogenetic shifts in migration in the North Atlantic.

Natal homing and connectivity in Atlantic bluefin tuna populations.

Atlantic bluefin tuna populations are in steep decline, and an improved understanding of connectivity between individuals from eastern (Mediterranean Sea) and western (Gulf of Mexico) spawning areas is needed to manage remaining fisheries. Chemical signatures in the otoliths of yearlings from regional nurseries were distinct and served as natural tags to assess natal homing and mixing. Adults showed high rates of natal homing to both eastern and western spawning areas. Trans-Atlantic movement (east to west) was significant and size-dependent, with individuals of Mediterranean origin mixing with the western population in the U.S. Atlantic. The largest (oldest) bluefin tuna collected near the northern extent of their range in North American waters were almost exclusively of western origin, indicating that this region represents critical habitat for the western population.