Basking sharks (Cetorhinus maximus) schooling in the southern Gulf of Maine

Schools of up to 50 basking sharks (Cetorhinus maximus) were observed on the warm side of a persistent thermal front in the southern Gulf of Maine from early September through mid‐October 2002. Aerial photographs showed a variety of schooling patterns, including echelon, cartwheel and milling formations. While the function of these aggregations remains unknown, they may represent group courtship behaviour.     

Modelling the transport of lobster (Homarus americanus) larvae and postlarvae in the Gulf of Maine

We used a coupled physical–biological model to examine potential distances between hatching and settlement locations for lobsters in the Gulf of Maine. The physical model is based on a finite-element mesh and climatological averages of the seasonally evolving temperature and density fields. Larval trajectories from coastal and offshore hatching sites (21–224 m deep) were calculated for early, middle and late-season hatching by coupling temperature-dependent development rates and depth (the biological model) to the circulation. Model results showed large spatial differences in larval development times (from 18 to 38 days) and distances transported (19–280 km) for the early hatch. Development time and transport decreased markedly by mid-season at most sites, but strong spatial differences persisted. The eastern Maine coast appears to experience stronger removal and less resupply of larvae than other regions, consistent with observed lower recruitment. Inverse solutions of the model for larvae arriving in mid-coastal Maine indicate that they originate from a broad section of the eastern coast 'upstream', with those nearest the shoreline generally travelling the shortest distances. The postlarval stage is neustonic (living near the surface), and a simple inverse model demonstrates that a diurnal coastal sea breeze can contribute substantially to inshore movement during this final planktonic stage. Thus, offshore reproduction may be linked to inshore recruitment.     

Spatiotemporal variability in the phenology of the initial intra‐annual molt of American lobster (Homarus americanus Milne Edwards, 1837) and its relationship with bottom temperatures in a changing Gulf of Maine

This study uses the Maine Department of Marine Resources Lobster Sea Sampling data (2000–2016) and logistic models to develop the first time series for the timing and suddenness of onset of the initial intra‐annual molt of American lobster in the Gulf of Maine (GoM), an annual fishery recruitment event crucial to fishermen. Data from three GoM regions (eastern, central, and western coastal Maine) were further divided by sex and estimated maturity of sampled lobsters for analysis. We found differences in the patterns of initial molt timing and suddenness between the regions, sexes, and stages of maturity. Using the Northeast Coastal Ocean Forecasting System hindcast temperatures, seasonal accumulated degrees above 5°C were used to describe the thermal history for each region at ocean depths of about 5 and 110 m. These temperature metrics were used in generalized linear models to investigate the potential effects of seasonal temperatures on the initial molt season. Results showed that initial intra‐annual molting of lobsters was variable from 2000 to 2016, with periods of both earlier and more sudden molts and later and more protracted molts. Warmer temperatures, specifically inshore temperatures, were generally associated with an earlier molt, but without complete uniformity in the direction and magnitude across seasons, regions, and lobster demographics. We also discuss why developing molt time series and quantifying the connection to the bottom temperatures are necessary and emphasized why existing monitoring programs and the applied quantification techniques herein make this relationship difficult to quantify.     

Bluefin tuna (Thunnus thynnus) distribution in relation to sea surface temperature fronts in the Gulf of Maine (1994–96)

Fishery‐linked aerial surveys for bluefin tuna (Thunnus thynnus) were conducted in the Gulf of Maine (GOM) from July through October, 1994–96. Each year, from 507 to 890 surface schools were detected and their locations examined in relation to oceanographic conditions. Correlations between bluefin tuna presence and environmental variables were explored for sea surface temperature (SST), distance to a SST front, frontal density (relative density of all SST fronts seen in a given 1 km area for 2 weeks prior to each tuna sighting), and bottom depth and slope. Mean SST associated with bluefin schools was 18.1°C (±2.8). Schools were located at a mean distance of 19.7 km (±19.6) from SST fronts, and in water masses with an average frontal density of 28.2 m km^?2 (±35.7). Mean bottom depth of detected schools was 139.0 m (±70.3), and mean bottom slope was 0.7% rise (±0.7). A binomial generalized linear model fit to these variables indicated that bluefin are seen closer to fronts than locations in which no tuna were seen. Using simple and partial Mantel tests, we investigated the spatial correlation between bluefin tuna presence and the environmental variables, controlling for spatial autocorrelation. For each day that schools were sighted, we performed 24 Mantel tests, on a combination of response and predictor variables. The spatial relationship between bluefin tuna and SST fronts was inconsistent. Our analysis identified significant spatial structure in the bluefin school locations that had no significant correlation with any of the measured environmental features, suggesting that other untested features, such as prey density, may be important predictors of bluefin distribution in the GOM.     

Processes controlling retention of spring‐spawned Atlantic cod (Gadus morhua) in the western Gulf of Maine and their relationship to an index of recruitment success

Atlantic cod, Gadus morhua, harvested in US waters are currently managed as a Gulf of Maine stock and as a stock comprising Georges Bank and southern New England populations. Over the past two and a half decades, success of age‐1 recruitment to the Gulf of Maine stock has varied by more than an order of magnitude. To investigate the hypothesis that this variation is related to variation in the transport of larval cod to nursery areas, we carried out model simulations of the movement of planktonic eggs and larvae spawned within the western Gulf of Maine during spring spawning events of 1995–2005. Results indicate that the retention of spring‐spawned cod, and their transport to areas suitable for early stage juvenile development, is strongly dependent on local wind conditions. Larval cod retention is favored during times of downwelling‐favorable winds and is least likely during times of upwelling‐favorable winds, during which buoyant eggs and early stage larvae tend to be advected offshore to the Western Maine Coastal Current and subsequently carried out of the Gulf of Maine. Model results also indicate that diel vertical migration of later stage larvae enhances the likelihood of retention within the western Gulf of Maine. Consistent with model results is a strong correlation between age‐1 recruitment success to the Gulf of Maine cod stock and the mean northward wind velocity measured in Massachusetts Bay during May. Based on these findings, we propose a wind index for strong recruitment success of age‐1 cod to the Gulf of Maine stock.     

Growth and mortality of cultured bay scallops in the Damariscotta River, Maine (USA)

Growth and mortality of hatchery-reared bay scallops, Argopecten irradians, were determined over an 18 month period (July 1994-December 1995) at two locations in the Damariscotta River, Maine, north of its natural distribution. Growth occurred primarily between May and October when mean water temperature exceeded 10 °C. Final mean shell height (56.3 mm) was significantly greater (p 0.0005) at the upriver site than at the downriver site in association with a higher mean water temperature. Cumulative mortality (43-49%) over this period occurred primarily over the winter months, in conjunction with mean water temperatures below 5 °C. Although it is possible to rear bay scallops to commercial size in Maine, the short growing season, high winter mortality, and costs associated with over-wintering and fouling will likely deter commercial culture.     

Evaluating spatio‐temporal variability in the habitat quality of Atlantic cod (Gadus morhua) in the Gulf of Maine

Reduced abundance and contracted spatial distribution of Atlantic cod, Gadus morhua, in the Gulf of Maine (GOM) may indicate large spatio‐temporal variation in their habitat quality. Season‐specific Habitat Suitability Index (HSI) models were developed to quantify such variation in the offshore GOM management area. Data used were non‐zero cod catch rates with calibrations from the Northeast Fisheries Science Center (NEFSC) spring and fall bottom trawl surveys over the period 1982–2013 and key physical environmental variables including depth, bottom temperature, bottom salinity and sediment types. Significant declines were found in the average HSI across the study area in the springs of early 2000s and 2010s. These low average HSI values coincide with reduced age‐1 recruitment of GOM cod stock after the mid‐1990s. Moreover, the western coastal areas of the GOM generally exhibited higher average HSI values than the eastern coastal areas, whereas the offshore areas always had the lowest average HSI. Relatively higher cod survey catch rates in the western GOM may imply positive influences of environmental controls on the distribution of GOM cod.     

Assessing the green sea urchin (Strongylocentrotus drobachiensis) stock in Maine, USA

The green sea urchin (Strongylocentrotus drobachiensis) fishery is of great importance to Maine’s economy. The fishery took off in the late 1980s as a result of expanding export markets, but has experienced substantial decline in landings since 1992 because of large decreases in urchin stock abundance. Fishery-independent surveys have not been conducted, and no formal stock assessment has been done prior to this study. Using the data collected from the fishery and urchin life history parameters derived from scientific studies, we conducted a formal stock assessment for the urchin stock. A stochastic observation-error length-structured model is used to describe the dynamics of the sea urchin population. A robust Bayesian approach is used for estimating fishery parameters because of concerns of possible outliers in fisheries data and mis-specification of priors. This study shows that the current stock is only 10% of the virgin stock biomass and that the exploitation rate is close to 40% suggesting that a large reduction in exploitation rate is necessary.     

Allozyme genetic analysis in hatchery strains and wild blue shrimp, Penaeus (Litopenaeus) stylirostris (Stimpson), from the Gulf of California

We conducted a population genetic analysis of Penaeus stylirostris to describe the genetic variability in two wild samples (Guaymas and Peñasco) and in two cultivated strains (A and B), and to determine the relationship among samples. Seventeen enzymatic systems and general proteins were used to visualize 31 loci. Of the 31 loci, 11 polymorphic loci (35%) were detected in all the organisms, with an average heterozygosity of 0.10 and an average of 1.8 per locus. There were no significant differences among the four samples of observed heterozygosity or percentage of polymorphic loci. According to Hardy–Weinberg, both cultivated strains showed disequilibrium in more loci than the wild samples. Guaymas showed linkage disequilibrium in five pairs of polymorphic loci, Peñasco samples in two pairs, A in seven pairs, and B in nine pairs. Peñasco and both cultivated lineages showed evidence of recent reductions in their effective populations size. Based on the comparisons of F_st values, and the distribution of the allelic frequencies, the cultivated strains were significantly different from the two wild populations. There is evidence that strain B originated from A and the discrepancies between the strains are probably the result of genetic drift and a founder effect.     

Modeled transport of winter flounder larvae spawned in coastal waters of the Gulf of Maine

Winter flounder, Pseudopleuronectes americanus, from southern New England and the Gulf of Maine were historically considered to be obligate estuarine spawners. However, recent experiments and observations document that winter flounder in the Gulf of Maine also utilize coastal waters for spawning. An individual‐based modeling approach was used to investigate the transport of winter flounder larvae from three hypothesized coastal spawning grounds in the Gulf of Maine. Transport success rates were greatest for larvae released from Ipswich Bay, intermediate for Stellwagen Bank and least successful for those released from Jeffreys Ledge. There was substantial interannual variability in larval transport and geographic patterns of potential connectivity. Furthermore, the date of spawning had an important influence on transport success. Model results suggest that certain coastal spawning grounds used by winter flounder may serve as an important source of larvae to estuaries and nearshore nursery areas. The potential influx of coastal spawned larvae could have implications for the resilience, productivity and gene flow in local populations. Model results provide further support for the conclusion that winter flounder in the Gulf of Maine may not be solely dependent upon estuaries for spawning. Results also suggest that coastal spawning groups should be considered explicitly in the management of winter flounder, and protected under Essential Fish Habitat regulations.     

Larval lobster (Homarus americanus) distribution and drift in the vicinity of the Gulf of Maine offshore banks and their probable origins

Surveys for lobster larvae in offshore waters of the north‐eastern Gulf of Maine in 1983, 1987 and 1989 confirm that local hatching occurs mainly at depths <100 m over the banks, including Georges and Browns Banks. Detailed studies in the vicinity of Georges Bank in late July of both 1987 and 1989 indicate that the first and second moult stages were located primarily over the bank whereas stages III and IV lobster were collected both over and off the bank. At times stage IV lobster were more abundant off the bank than over it. The condition of stage III and IV lobster, as measured by a lipid index, was better off than over Georges Bank in 1988 and 1989 indicating a possible physiological advantage to being off the bank. In addition, the higher surface temperatures off Georges Bank would shorten larval development time to settlement. To determine the probable hatch sites of stage IV lobster collected off of Browns Bank in 1983 and off of Georges in 1987 and 1989, a 3‐D circulation model of the Gulf of Maine was used to simulate larval lobster drift backwards in time. In all cases, areas off Cape Cod, MA, and off Penobscot Bay, ME were suggested as the source of the larvae, although most of the larval trajectories never reached these near‐shore waters that are well‐known, larval hatching areas. The model‐projected larval release times match most closely the observed inshore hatch off Massachusetts but model uncertainties mean that coastal Maine cannot be ruled out as a source. Georges Bank is also a potential source because the present model does not take into account short‐term wind events, off‐bank eddy transport or the possibility of directed off‐bank larval swimming. Examination of weather records prior to and during our 1988 and 1989 sampling periods indicates that winds were not of sufficient intensity and duration to induce larval transport off Georges Bank. The shedding of eddies from the northern flank of Georges Bank into the Gulf of Maine are a relatively common phenomenon during summer but not enough is known about them to evaluate their contribution to possible cross‐bank transport of lobster larvae. Directed larval swimming is another possible source for the stage IV lobster found near Georges Bank. Plankton distributions across the northern frontal zone of Georges Bank in 1988 were used as proxies for the scarce larval lobsters. The more surface distribution of the microplankton, in particular, supports the possibility that wind and eddy events may be important in the transport of stage III and IV lobsters off of Georges Bank. Further studies are needed to evaluate these possible additional sources of advanced stage lobster larvae found off of the offshore banks.     

Multidecadal otolith growth histories for red and gray snapper (Lutjanus spp.) in the northern Gulf of Mexico,USA

Dendrochronology (tree‐ring analysis) techniques were applied to develop chronologies from the annual growth‐increment widths of red snapper (Lutjanus campechanus) and gray snapper (Lutjanus griseus) otoliths sampled from the northern Gulf of Mexico, USA. Growth increment widths showed considerable synchrony within and across species, indicating that some component of environmental variability influenced growth. The final, exactly dated red snapper chronology continuously spanned 1975 through 2003, while the gray snapper chronology continuously spanned 1975 through 2006. To determine baseline climate‐growth relationships, chronologies were compared to monthly averages of sea surface temperatures, U winds (west to east), V winds (south to north), and Mississippi River discharge. The gray snapper chronology significantly (P < 0.01) correlated with winds and temperature in March and April, while the red snapper chronology correlated with winds in March. Principal components regression including springtime winds and temperature accounted for 28 and 52% of the variance in the red and gray snapper chronologies, respectively. These results indicate that snapper growth was favored by warm sea surface temperatures and onshore winds from the southeast to the northwest in March and April. Overall, this study provides preliminary, baseline information regarding the association between climate and growth for these commercially important snapper species.     

The interaction of infectious salmon anaemia virus (ISAV) with the blue mussel,Mytilus edulis

Integrated multi‐trophic aquaculture (IMTA) is an alternative approach to mono‐culture aquaculture that reduces environmental impacts of commercial aquaculture systems by combining the cultivation of fed species with extractive species. Shellfish play a critical role in IMTA systems by filter‐feeding particulate‐bound organic nutrients. They may also increase or decrease disease risk on farms by serving as reservoirs or barriers for important finfish pathogens such as infectious salmon anaemia virus (ISAV). This study aimed to optimize culture and molecular assays in shellfish tissues and to determine the fate of ISAV in mussels, Mytilus edulis. To determine detection limits, qRT‐PCR and culture assays in both CHSE‐ and ASK cells were optimized in ISAV‐inoculated mussel tissue homogenates. Both qRT‐PCR and culture assays performed in ASK cells had comparable detection limits of 10^2.8 TCID₅₀ mL^?1. The ISAV RNA genome was consistently detected in digestive gland tissue of ISAV‐exposed mussels. Viable ISAV was not detected in mussel tissues by culture analysis in CHSE‐ and ASK cells. The fact that qRT‐PCR analysis resulted in positive cycle threshold (CT) values that corresponded to the detectable range of ISAV in ASK culture assays suggests that little to no viable ISAV particles are present in the mussel tissues.     

It’s about time: A synthesis of changing phenology in the Gulf of Maine ecosystem

The timing of recurring biological and seasonal environmental events is changing on a global scale relative to temperature and other climate drivers. This study considers the Gulf of Maine ecosystem, a region of high social and ecological importance in the Northwest Atlantic Ocean and synthesizes current knowledge of (a) key seasonal processes, patterns, and events; (b) direct evidence for shifts in timing; (c) implications of phenological responses for linked ecological‐human systems; and (d) potential phenology‐focused adaptation strategies and actions. Twenty studies demonstrated shifts in timing of regional marine organisms and seasonal environmental events. The most common response was earlier timing, observed in spring onset, spring and winter hydrology, zooplankton abundance, occurrence of several larval fishes, and diadromous fish migrations. Later timing was documented for fall onset, reproduction and fledging in Atlantic puffins, spring and fall phytoplankton blooms, and occurrence of additional larval fishes. Changes in event duration generally increased and were detected in zooplankton peak abundance, early life history periods of macro‐invertebrates, and lobster fishery landings. Reduced duration was observed in winter–spring ice‐affected stream flows. Two studies projected phenological changes, both finding diapause duration would decrease in zooplankton under future climate scenarios. Phenological responses were species‐specific and varied depending on the environmental driver, spatial, and temporal scales evaluated. Overall, a wide range of baseline phenology and relevant modeling studies exist, yet surprisingly few document long‐term shifts. Results reveal a need for increased emphasis on phenological shifts in the Gulf of Maine and identify opportunities for future research and consideration of phenological changes in adaptation efforts.     

An adjoint data assimilation approach to diagnosis of physical and biological controls on Pseudocalanus spp. in the Gulf of Maine–Georges Bank region

The underlying scientific objective here is to determine the mechanisms that control seasonal variations in the abundance of Pseudocalanus spp. in the Georges Bank–Gulf of Maine region. It is postulated that the observed distributions result from the interaction of the population dynamics with the climatological circulation. The problem is posed mathematically as a 2-D advection–diffusion–reaction equation for a scalar variable. Given an initial distribution of animals, we seek the population dynamics source term R ( x , y ) such that integration of the forward model will result in predictions3 that minimize the sum of squares of differences with observed concentrations at a later time. An adjoint data assimilation technique has been designed for these purposes.
This approach has been used to invert for the population dynamics associated with the transition between bimonthly (i.e. for 2 months) climatological Pseudocalanus spp. distributions derived from MARMAP data. Vertically averaged velocity and diffusivity fields diagnosed from hydrodynamical simulations of the climatological flow are specified. Solutions converge rapidly, and the procedure reduces the cost function by an order of magnitude within 50 iterations. The resulting population dynamics vary considerably in space and time, as does the balance between local tendency, physical transport and biological source terms. Generally speaking, the patterns in population dynamics are not inconsistent with current knowledge concerning potential controls such as predation and food limitation. Analysis of the solutions indicates that the Pseudocalanus spp. population centres located in the western Gulf of Maine and on Georges Bank may be self-sustaining, in contrast to prior studies which characterize the former as a source region for the latter.     

Cadmium,lead and metallothionein contents in cultivated mussels (Mytilus galloprovincialis) from the Gulf of Naples (Southern Italy)

The concentration of cadmium and lead ions was determined in digestive gland and mantle of mussels (Mytilus galloprovincialis) from five aquaculture stations located in the Gulf of Naples (Southern Italy, Tyrrhenian Sea). Metallothionein (MT) levels in the same tissues were also evaluated. This gulf represents a greatly urbanized area, characterized by an important commercial port and past industrial activities, terminated in the last decade of the 20th century. The results were compared with those obtained from mussels cultivated in a more pristine neighbouring area. Data demonstrate that the amount of both metals found in mussels harvested at the highly anthropogenic sites in the Gulf of Naples was comparable with that found in mussels from the more naturalistic site. The content of both cadmium and lead was not related with that of MT. Indeed, the amount of both metals in the digestive gland was greater than in the mantle, whereas the MT preferentially accumulated in the mantle. Possibly, the MT content in mantle was associated to the physiological function of the tissue, rather than heavy metals exposure.     

Effects of seed size and density on growth, tissue-to-shell ratio and survival of cultivated mussels (Mytilus edulis) in Prince Edward Island, Canada

In suspended aquaculture, the size of mussel seed and the density at which these mussels are placed in socks are two important factors affecting mussel production. However, the effect of these two factors has yet to be examined simultaneously. We conducted two large-scale field experiments to assess the effect of seed size and initial density on growth, tissue-to-shell ratio and survival of blue mussels (Mytilus edulis) grown on longlines. Experimental mussel sites were set up during the autumn of two consecutive years in two bays on the north shore of Prince Edward Island, Canada. At each site, socks containing combinations of three seed sizes (small, medium and large) at two initial densities (low and high) were deployed and monitored over time. Small seed grew faster than large seed and often reached commercial size in the same time period. Furthermore, initial density generally did not affect shell growth. During the pre-spawning period following the autumn deployment, seed at low initial density had a higher tissue-to-shell ratio than seed at high initial density, especially for large seed. This effect of initial density disappeared in early summer. Early in the experiments (< about 10 months), mussel survival on socks was affected by seed size only: small seed generally had lower survival rates than larger seed. Late in the experiments (> 10 months), there appeared to be an interaction between seed size and initial density on mussel survival: survival of small seed generally decreased with increasing initial density, while survival of large seed was not affected by initial density. Since large seed tended to be packed at lower initial density than small seed, lack of crowding was likely responsible for the absence of density-dependent loss. Our study thus assessed different seeding strategies, and results were generally consistent across years and sites.     

Use of satellite data to identify critical periods for early life survival of northern shrimp in the Gulf of Maine

The northern shrimp Pandalus borealis is at its southern limit in the Gulf of Maine (GOM), and recruitment success is higher in years with relatively cool water temperature. However, the mechanisms for the temperature effect are not clear. We used rolling window analysis of daily satellite data to identify critical periods for early life survival of the 1998–2012 northern shrimp year‐classes and to investigate the importance of the phenology of the hatch and bloom. Survival was negatively correlated with sea surface temperature (SST) during a 6‐week period around the time of larval emergence (late winter) and during a 4‐week period in late summer when SST and stratification reached annual maxima. Survival was negatively correlated with chlorophyll‐a concentration (chl‐a) during two 5‐week periods centered approximately a month before the hatch midpoint and around the time of settlement to the benthos. A small‐magnitude winter bloom occurred around the time of the hatch in many years, but our results did not reveal a link between survival and bloom‐hatch phenology. The timing of winter and spring blooms were correlated with SST during the preceding 10 months. A survival model including SST and chl‐a during the critical periods explained 73% of the variance in survival. Summer SST increased significantly during the study period; the other critical variables showed no trend. The rolling windows approach revealed sensitive periods in early life history that may not have otherwise been hypothesized, providing a foundation for research towards a greater understanding of processes affecting recruitment.