Interannual variability in growth of larval and juvenile walleye pollock Theragra chalcogramma in the western Gulf of Alaska, 1983–91

Interannual variability in growth of larval walleye pollock Theragra chalcogramma was examined from 1983 to 1991 and of juveniles from 1985 to 1990. ANCOVA was used to assess differences in population growth rates, and an alternate method was developed to examine variations between annual length-at-age data and average 'expected' values over different age groupings. For larvae, the years 1986, 1987, 1989 and 1990 had higher than average length-at-age, and 1988 and 1991 had lower than average values. Relationships between growth and SST and larval density were not clear. A tentative relationship between copepod nauplii abundance and larval length-at-age was noted. The consequence of larval growth to larval mortality, late larval abundance or recruitment was not clear. We conclude that larval mortality rates are highly variable and tend to mask effects of moderate variability in growth on later abundance. For juveniles, 1987 had significantly lower than average length-at-age and 1988 had higher than average values. Although there are few years of data, they tend to support the importance of juvenile growth in the recruitment process. Conditions for the large 1988 year class are documented and discussed, including warm SST, calm winds, relatively low larval growth rates, low abundances of potential predators on larvae, low larval mortality rates, and high juvenile growth rates.     

Density‐ and size‐dependent mortality in fish early life stages

The importance of survival and growth variations early in life for population dynamics depends on the degrees of compensatory density dependence and size dependence in survival at later life stages. Quantifying density‐ and size‐dependent mortality at different juvenile stages is therefore important to understand and potentially predict the recruitment to the population. We applied a statistical state‐space modelling approach to analyse time series of abundance and mean body size of larval and juvenile fish. The focus was to identify the importance of abundance and body size for growth and survival through successive larval and juvenile age intervals, and to quantify how the dynamics propagate through the early life to influence recruitment. We thus identified both relevant ages and mechanisms (i.e. density dependence and size dependence in survival and growth) linking recruitment variability to early life dynamics. The analysis was conducted on six economically and ecologically important fish populations from cold temperate and sub‐arctic marine ecosystems. Our results underscore the importance of size for survival early in life. The comparative analysis suggests that size‐dependent mortality and density‐dependent growth frequently occur at a transition from pelagic to demersal habitats, which may be linked to competition for suitable habitat. The generality of this hypothesis warrants testing in future research.     

Spatial distribution and overlap between ichthyoplankton and pelagic fish and squids on the southern flank of Georges Bank

We conducted larval and adult fish surveys on the southern flank of Georges Bank during the spring of two years (1990 and 1995) with contrasting physical conditions. We employed canonical correspondence analysis (CCA) to examine the relationships between physical variables and the spatial distribution of pelagic fish and ichthyoplankton. Surface temperature bottom temperature, and vertical stratification were significant factors affecting larval fish distributions, and there were groups of species with similar responses to these variables. There were also consistent relationships between physical variables and pelagic fish and squid abundances and spatial distributions. Pelagic fish and ichthyoplankton with similar responses to hydrographic variables had high spatial overlap, and variation in hydrographic regimes modulated the strength of this interaction. Pelagic fish and squids are potentially important predators of both larval and juvenile fish on Georges Bank. Hydrographic structure modulates the degree of spatial overlap and therefore likely influences the strength of predator–prey interactions.     

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.     

Do albacore exert top‐down pressure on northern anchovy? Estimating anchovy mortality as a result of predation by juvenile north pacific albacore in the California current system

Quantifying the mortality of marine fishes is important for understanding spawner–recruit relationships, predicting year‐class strength, and improving fishery stock assessment models. There is increasing evidence that pelagic predators can exert a top‐down influence on prey, especially during critical early life‐history stages. The objective of this study was to quantify predation by North Pacific albacore on Northern anchovy in the California current system (CCS). I estimated the abundance of juvenile albacore in the CCS from 1966–2005 using stock assessment models and spatially explicit catch‐per‐unit‐effort time series. Anchovy abundance (1966–93), both recruitment and total biomass, was obtained from a stock assessment model. Annual rates of anchovy consumption by albacore were calculated using diet studies of albacore in the CCS, an age‐structured bioenergetics model, and regional estimates of albacore abundance. The range of estimates was large: albacore may remove from less than 1% to over 17% of anchovy pre‐recruitment biomass annually. Relationships between predation and recruitment biomass were consistent with expectations from top‐down effects, but further study is required. This is the first attempt to quantify a specific source of mortality on anchovy recruits and to demonstrate potential top‐down effects of predation on anchovy.     

Modeling the effects of temperature on the survival and growth of North Sea cod (Gadus morhua) through the first year of life

Temperature and body size are widely agreed to be the primary factors influencing vital rates (e.g., growth, mortality) in marine fishes. We created a biophysical individual‐based model which included the effects of body size and temperature on development, growth and mortality rates of eggs, larvae and juveniles of Atlantic cod (Gadus morhua L.) in the North Sea. Temperature‐dependent mortality rates in our model were based on the consumption rate of predators of cod early‐life stages. The model predicted 35%, 53% and 12% of the total mortality to occur during the egg, larval and juvenile stages, respectively. A comparison of modeled and observed body size suggested that the growth of survivors through their first year of life is high and close to the growth rates in ad libitum feeding laboratory experiments. Furthermore, our model indicates that experiencing warmer temperatures during early life only benefits young cod (or theoretically any organism) if a high ratio exists between the temperature coefficients for the rate of growth and the rate of mortality. During the egg stage of cod, any benefit of developing more rapidly at warmer temperatures is largely counteracted by temperature‐dependent increases in predation pressure. In contrast, juvenile (age‐0) cod experiences a higher cumulative mortality at warmer temperatures in the North Sea. Thus, our study adds a new aspect to the ‘growth–survival’ hypothesis: faster growth is not always profitable for early‐life stages particularly if it is caused by warmer temperatures.     

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.     

Interannual variation in quantitative relationships among egg production and densities of larvae and juveniles of the Japanese mantis shrimp <Emphasis Type="Italic">Oratosquilla oratoria</Emphasis> in Tokyo Bay,Japan

To explore which lifestages affect the stock size of young-of-the-year mantis shrimp Oratosquilla oratoria in Tokyo Bay, Japan, we investigated interannual variations in the quantitative relationships among egg production, larval density, and juvenile density. We collected adult females, larvae, and juveniles during monthly field surveys from 2004 to 2007. The interannual trend for the juvenile density index differed from those for egg production and larval density; although indices of both egg production and larval density were high in 2004 and 2007, the juvenile density index was high only in 2007, suggesting high mortality during the pelagic larval stage or the early phase of the postsettlement juvenile stage in 2004. We found that larval settlement started at the end of August and peaked in October, although larvae from the early spawning season (May–June) should have settled in August or earlier. Juveniles were found throughout the bay except in areas where bottom hypoxia occurred, suggesting that hypoxia restricts the spatial distribution of juveniles. Our results suggest that mortality during the early life history fluctuates among years, probably because of changes in environmental conditions in the bay, resulting in interannual variation in the stock size of young-of-the-year juvenile O. oratoria.     

Recruitment variability and oceanographic stability

We examine the hypothesis that recruitment is more variable in populations on isolated offshore banks than nearby shelf populations. Recruitment of cod (Gadus morhua) and American plaice (Hippoghssoides plates-soides) on Flemish Cap is more variable than in any comparable population. Recruitment of haddock (Melanogrammus aeglefinus) on Rockall Bank is also more variable than in surrounding populations. These results are confirmed both by estimates obtained by virtual population analysis and by research surveys. Recruitment of haddock and herring (Clupea harengus) on Georges Bank is also more variable than in surrounding populations; however, the results for two other groundfish populations, cod and yellowtake flounder (Limanda ferruginae), on Georges Bank are ambiguous. We conclude that marine fish populations on isolated banks are more variable than those on nearby shelf regions.     

Settlement and recruitment patterns of Mytilus galloprovincialis L. in the Ría de Ares-Betanzos (NW Spain) in the years 2004/2005

The present study explores the settlement and recruitment capacity of Mytilus galloprovincialis L. on artificial collectors in the Ría de Ares‐Betanzos (Galicia, NW Spain) in 2004 and 2005 following standard industrial techniques. Three locations in the ría (Arnela, Redes and Miranda) were selected to investigate larvae settlement after the main spawning event (July 2004/2005). Assessment of the recruitment capacity was performed in autumn (September 2004/2005) when mussel seed is usually gathered from the collector ropes and introduced into industrial cultivation at low densities. For both years, the highest settlement densities were recorded at the most seaward location, Miranda. Differences in settlement densities between locations are discussed in terms of the water circulation regime in the ría and the local hydrographic conditions. In 2004, statistical differences in post‐settlement mortality resulted in similar recruitment densities at Arnela and Miranda, which were higher than at Redes. In 2005, recruitment densities in Redes and Arnela could not be assessed because predators (Spondyliosoma cantharus L.) eliminated the settled population at these locations. Site‐specific differences in recruitment density may be attributed to the environmental limitations of each location as well as intra‐specific competition.     

Modeling larval Calanus finmarchicus on Georges Bank: time-varying mortality rates and a cannibalism hypothesis

Throughout the North Atlantic, the copepod Calanus finmarchicus dominates the zooplankton biomass, linking primary production and higher trophic levels. On Georges Bank, the peak abundance of larval (naupliar) stages occurs in March–April and represents a potential source of prey for cod and haddock larvae. Following this maximum, naupliar abundance declines dramatically, reaching a minimum in May and increasing again in June. Explaining the naupliar seasonal cycle is critical for predicting climate effects on C. finmarchicus dynamics, including whether environmental variability may lead to a mismatch with larval fish. Here, an age-within-stage population dynamics model is used to investigate the factors controlling the temporal variation of C. finmarchicus nauplii in three Georges Bank sub-regions. The model incorporates temperature- and food-dependent development and egg production, as well as female abundance derived from the US Global Ocean Ecosystem Dynamics (GLOBEC) program. Use of field-estimated constant mortality rates overestimates May abundances by as much as an order of magnitude. These data/model discrepancies can not be explained by temperature or food-limitation effects on physiological rates. Instead, accurate simulation requires use of time-varying early stage mortalities, which differ from published estimates in both magnitude and trend. These mortality rates are correlated with C. finmarchicus female abundance, implying cannibalism as a possible regulatory factor. Thus, the biological control of predation (including cannibalism) must be considered to predict the effects of climate on C. finmarchicus and associated larval fish populations.     

Potential of contemporary evolution to erode fishery benefits from marine reserves

Marine reserves are valued for their ecological role: protecting fish populations from overharvesting while, at the same time, potentially maintaining fisheries yields via recruitment effects (net export of pelagic eggs and larvae) and spillover (net export of post‐settled juveniles and mature fish) across reserve borders. Focussing on the spillover effect, we argue that when fitness of the protected individuals depends on the relative size of their home ranges compared to the reserve size, and home range size is a property of the individuals, rapid local adaptation might occur in favour of individuals with smaller home ranges. Individuals that avoid fishing mortality by spending most of the time inside the reserve limits (i.e. with smaller home ranges) will experience a fitness advantage, whereas individuals that move beyond boundaries (i.e. with larger home ranges) will increase their risk of being harvested by spillover fisheries. We use empirical data on Atlantic cod (Gadus morhua) home ranges within and around a coastal marine reserve in south Norway as a case to illustrate our idea. Broadly, we highlight how protection‐induced selection and contemporary evolution could fundamentally alter our perspective of marine reserve functioning and recommend quantifying behavioural variability and behavioural consistency of protected populations.     

Towards a more complete understanding of the life cycle of brown shrimp (Crangon crangon): modelling passive larvae and juvenile transport in combination with physically forced vertical juvenile migration

In this study we developed and utilized a complex model approach to investigate the impact of stage‐specific transport processes on the development and spatial distribution of brown shrimp (Crangon crangon) post‐larvae and juveniles in the German Bight. First, we focused on drift processes during the pelagic larval stage by coupling an individual‐based model for egg and larval development ‘off‐line’ to a 3D hydrodynamic model utilizing the Lagrangian method. Secondly, we investigated tidal‐induced transport processes after juvenile settlement. To determine the tidal cycle, the model coupling was accomplished ‘on‐line’ by resolving the individual‐based model and hydrodynamic model with the same time step. The vertical migration of juveniles, a prerequisite for the selective tidal stream transport (STST), was modelled as a sub‐grid scale physical process (balance of forces: gravitation, buoyancy, Stoke’s friction and dynamic uplift) and considered complex particle dynamics. We applied the model to test temperature and salinity cues as possible tidal indicators utilized by juvenile brown shrimp. Our results indicated that transport processes could significantly change the timing and spatial distribution of post‐larval abundance. We also showed that the small‐scale hydrodynamic forcing acting on the bodies of juvenile brown shrimps was sufficient to account for the vertical migration required to use STST. For both investigated tidal cues STST performing juvenile brown shrimp were transported on‐shore. A faster and more continuous STST was calculated for the salinity cue, resulting in larger abundances of brown shrimp in estuarine areas.     

Simulation of mackerel (Scomber scombrus) recruitment with an individual-based model and comparison with field data

An individual‐based model (IBM) for the simulation of year‐to‐year survival during the early life‐history stages of the north‐east Atlantic stock of mackerel (Scomber scombrus) was developed within the EU funded Shelf‐Edge Advection, Mortality and Recruitment (SEAMAR) programme. The IBM included transport, growth and survival and was used to track the passive movement of mackerel eggs, larvae and post‐larvae and determine their distribution and abundance after approximately 2 months of drift. One of the main outputs from the IBM, namely distributions and numbers of surviving post‐larvae, are compared with field data as recruit (age‐0/age‐1 juveniles) distribution and abundance for the years 1998, 1999 and 2000. The juvenile distributions show more inter‐annual and spatial variability than the modelled distributions of survivors; this may be due to the restriction of using the same initial egg distribution for all 3 yr of simulation. The IBM simulations indicate two main recruitment areas for the north‐east Atlantic stock of mackerel, these being Porcupine Bank and the south‐eastern Bay of Biscay. These areas correspond to areas of high juvenile catches, although the juveniles generally have a more widespread distribution than the model simulations. The best agreement between modelled data and field data for distribution (juveniles and model survivors) is for the year 1998. The juvenile catches in different representative nursery areas are totalled to give a field abundance index (FAI). This index is compared with a model survivor index (MSI) which is calculated from the total of survivors for the whole spawning season. The MSI compares favourably with the FAI for 1998 and 1999 but not for 2000; in this year, juvenile catches dropped sharply compared with the previous years but there was no equivalent drop in modelled survivors.     

Using a biophysical model to investigate connectivity between spawning grounds and nursery areas of King George whiting (Sillaginodes punctatus: Perciformes) in South Australia’s gulfs

Many demersal marine fish species depend on a dispersive larval stage that connects geographically discrete sub‐populations. Understanding connectivity between these sub‐populations is necessary to determine stock structure, which identifies the appropriate spatial scale for fishery management. Such connectivity is poorly understood for King George whiting (Sillaginodes punctatus; Perciformes) in South Australia's gulf system, even though spawning grounds and nursery areas are adequately defined. In response to declines in commercial catches and estimated biomass, this study aimed to determine the most important spawning grounds and nursery areas to recruitment, and the connectivity between them. A biophysical model was seeded with particles according to the distribution and density of eggs throughout the spawning area in 2017 and 2018. Despite inter‐annual differences in the origins of particles, dispersal pathways and predicted settlement areas remained consistent between years. Predicted settlement was generally highest to nursery areas only short distances from regional spawning grounds, consistent with previous hydrodynamic models. However, the model also predicted that spawning in one region could contribute to recruitment in an adjacent region later in the spawning season, which aligned with the breakdown of thermohaline fronts at the entrance of each gulf. The connectivity between spawning grounds and nursery areas predicted by the model is supported by spatio‐temporal patterns in the otolith chemistry of pre‐flexion larvae and settled juveniles. Consequently, the most parsimonious explanation is that the populations of King George whiting in South Australia's gulf system constitute a single, panmictic stock, which has implications for fishery management.     

Interannual variation in the coastal distribution of a juvenile gadid in the northeast Pacific Ocean: The relevance of wind and effect on recruitment

Drift of propagules occurs within many populations inhabiting flow fields. This affects the number of propagules that rejoin their source population (recruitment) and plays a role in adaptive spatial redistribution. We focus on the cause and consequence of interannual variation in geographic distribution of population density among five cohorts of young‐of‐the‐year (age‐0) juvenile walleye pollock Gadus chalcogrammus in the western Gulf of Alaska (GOA). The coastal GOA is a wind‐driven advective system. Walleye pollock spawn during spring and their eggs and larvae drift southwestward; by late summer, age‐0 juveniles are variously distributed over the shelf. We found that high population densities of age‐0 juveniles (ca. 6 months old) near the southwestward exit of the Alaska Coastal Current from the GOA corresponded with high abundance of larvae from the major spawning area upstream, but did not translate into high abundance at older ages. Further, offshore and upwelling‐favorable winds were associated with the high downstream abundance and presumed export. In contrast, downwelling‐favorable (northeasterly) wind during and shortly after spawning (April–May) was associated with high recruitment at age 1. Finally, we found that recruitment also increased with apparent retention of age‐0 juveniles in favorable habitat upstream near the main spawning area. We hypothesize that wind‐related retention in superior upstream habitat favors recruitment. Our results argue for including wind‐driven transport in future walleye pollock recruitment models. We encourage more work on the juvenile stage of marine fishes aimed at understanding how transport and species‐specific habitat suitability interact to affect population response to large‐scale forcing.     

Optimum temperature and food-limited growth of larval Atlantic cod (Gadus morhua) and haddock (Melanogrammus aeglefinus) on Georges Bank

We estimated recent growth of Atlantic cod (Gadus morhua) and haddock (Melanogrammus aeglefinus) larvae collected on the southern flank of Georges Bank in May 1992–94 from the ratio of RNA to DNA (R/D) and water temperature. Growth of both species increased with water temperature to about 7°C and then decreased. The highest growth rates were observed in May 1993 at water temperatures around 7°C. These data confirm an earlier observation of comparable temperature optima for growth of Atlantic cod and haddock larvae in the north‐west Atlantic. Comparisons of field growth rates and temperature optima with data for larvae cultured at high temperatures and prey densities in the laboratory suggest that growth may have been food‐limited at higher temperatures on Georges Bank. Given that 7°C is the long‐term mean water temperature on the southern flank in May and that climate models predict a possible 2–4°C rise in water temperatures for the western North Atlantic, our findings point to a possible adverse effect of global warming on Atlantic cod and haddock.     

Scleractinian coral communities of the inner Seychelles 10 years after the 1998 mortality event

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Comparing the roles of Pacific halibut and arrowtooth flounder within the Gulf of Alaska ecosystem and fishing economy

The fishing industry of the western and central regions of the coastal Gulf of Alaska (CGoA) directly employs over 17,000 people and processes fish with a wholesale value of US$618 million annually. Pacific halibut (Hippoglossus stenolepis) are a valued groundfish species because of the high quality of their flesh. In contrast, arrowtooth flounder (Atheresthes stomias) are much more abundant but of low value because their flesh degrades upon heating. Both are high trophic level predators but play different roles in the ecosystem because of differences in abundance and diet. Using an end‐to‐end ecosystem model, we evaluate the impact of alternate levels of fishing effort and large‐scale changes in oceanographic conditions upon both species, the ecosystem, and the fishing economy. Reduction of longline efforts to reduce Pacific halibut mortality led to reduction in total value of all CGoA landings but increase in value landed by sport fisheries, trawl fleets, and fish pot vessels as they exploit a greater share of available halibut, sablefish, and Pacific cod. Increased trawl effort to raise arrowtooth flounder mortality led to increase in total value of all landings but large reductions in value landed by longline, jig, fish pot, and sport fleets with greater competition for available Pacific cod, halibut, and sablefish. Oceanographic conditions that enhance pelagic food chains at the expense of benthic food chains negatively impact groundfish in general, though Pacific halibut and arrowtooth flounder are resilient to these effects because of the high importance of pelagic fish in their diets.