Changes in the spawning habitat of two small pelagic fish in the Northwestern Mediterranean

We investigated the effects of three sea surface oceanographic variables (temperature, salinity, and chlorophyll a) on the abundance of eggs and larvae of two summer‐spawning species in the NW Mediterranean Sea, the anchovy Engraulis encrasicolus and the round sardinella Sardinella aurita, based on data from ichthyoplankton surveys carried out in the 1980s, 2000s, and 2010s. The environmental data showed an increase in seawater temperature and salinity along time, coupled with a decrease in chlorophyll a (proxy for primary production). These long‐term directional changes in environmental conditions helped explain the important reduction observed in the abundance of eggs and larvae of anchovy, as well as shrinking of spawning habitat in this species. At the same time, the probability of occurrence of round sardinella has increased from practically zero in the 1980s to probabilities near 1 along the coastal area of the study region in the two decades of the 21st century. Given that the trends observed in the environmental variables along the three decades of study are expected to continue during the 21st century, as a consequence of climate change, the spawning habitat of anchovy is expected to continue decreasing, while round sardinella habitat can expand. Considering that anchovy is of high commercial importance in NW Mediterranean fisheries, while round sardinella has very low commercial interest, our results show that the viability of small pelagic fisheries in the area may be compromised.     

Alternating dominance of postlarval sardine and anchovy caught by coastal fishery in relation to the Kuroshio meander in the Enshu-nada Sea

In the mid 1970s, the fishery catch of postlarval Japanese anchovy (Engraulis japonica) in a shelf region of the Enshu‐nada Sea, off the central Pacific coast of Japan, started to decline corresponding to a rapid increase of postlarval sardine (Sardinops melanostictus). In late 1980s, sardine started to decline, and it was replaced by anchovy in the 1990s. This alternating dominance of postlarval sardine and anchovy corresponded to the alternation in egg abundance of these two species in the spawning habitat of this sea. It was also noteworthy that during the period of sardine decline, sardine spawning occurred in April–May, a delay of two months compared with spawning in the late 1970s. The implication of oceanographic changes in the spawning habitat for the alternating dominance of sardine and anchovy eggs was explored using time‐series data obtained in 1975–1998, focusing on the effect of the Kuroshio meander. Large meanders of the Kuroshio may have enhanced the onshore intrusion of the warm water into the shelf region and contributed to an increase in temperature in the spawning habitat. This might favour sardine, because its egg abundance in the shelf region was more dependent on the temperature in early spring than was that of anchovy. In addition, enhanced onshore intrusion could contribute to transport of sardine larvae from upstream spawning grounds of the Kuroshio region. On the other hand, anchovy egg abundance was more closely related to lower transparency at the shelf edge, which may indicate the prevalence and prolonged residence of the coastal water, and therefore higher food availability, frequently accompanying non‐meandering Kuroshio. The expansion/shrinkage of the spawning habitat of sardine and anchovy in the shelf region, apparently responding to the change in the Kuroshio, possibly makes the alternation in dominance of postlarval sardine and anchovy most prominent in the Enshu‐nada Sea, in combination with changes in the abundance of spawning adults, which occurred almost simultaneously in the overall Kuroshio region. The implication of this rather regional feature for the alternating dominance of sardine and anchovy populations on a larger spatial scale is also discussed.     

Modeling environmental,temporal and spatial effects on twaite shad (Alosa fallax) by‐catches in the central Mediterranean Sea

Modeling the relationships between environmental factors and the distribution at sea of species of conservation interest can be useful in predicting their occurrence from a local to a regional scale. This information is essential for planning management and conservation initiatives. In this study, generalized additive models (GAMs) were applied to investigate the influence of environmental, temporal and spatial variables on the catch rates of the twaite shad Alosa fallax (Lacepède) by the pelagic trawl fishery in the north‐central Adriatic Sea. Presence/absence and abundance [catch per unit effort (CPUE)] data between 2006 and 2012 were separately modeled, and the two models were then validated using a test data set. The most important factor influencing the presence and abundance of adult twaite shads was the spatial predictor (latitude × longitude). Two areas of major shads aggregations were observed, the most important of which being located near the estuaries of three main river systems of northern Italy. The twaite shad presence was also significantly affected by season, the largest and lowest occurrences being observed in autumn and spring, respectively. Among the environmental variables tested, only sea surface temperature was included in both models. Alosa fallax showed a wide thermal tolerance (6–27°C) with preference for temperature around 23°C. The model developed from the abundance data showed a moderate predictive power, whereas the accuracy of the presence/absence model was rather low. Some conclusions on the ecological requirements of A. fallax at sea arising from this study are useful to orient future monitoring and research programs and to develop effective conservation actions.     

Interannual decrease in condition factor of the western sand lance Ammodytes japonicus in Japan in the last decade: Evidence for food‐limited decline of the catch

Annual catch of the western sand lance Ammodytes japonicus in the eastern Seto Inland Sea, Japan, has shown a decreasing trend since the 1990s. To examine whether food shortage was the main cause for the catch decrease, we investigated the condition factor of the age‐0 fish at the beginning of the estivation period (late July) in Harima‐Nada, eastern Seto Inland Sea, for 10 years from 2008 to 2017. The mean abundance of copepods as food for the age‐0 fish during the fish growth period (from February to June) around the estivation area was also determined. The condition factor showed a significant decrease, and values for 2011 and later years were mostly lower than the known minimum threshold (4.2) for maturation. In the recent 4 years from 2013 to 2016, the mean copepod abundance was much lower than values for the other years. The condition factor showed a significant positive correlation with the copepod abundance. These results indicate that decline of western sand lance catch after 1990 was caused mainly by food shortage.     

Environmental effects on the spatio‐temporal patterns of abundance and distribution of Sardina pilchardus and sardinella off the Mauritanian coast (North‐West Africa)

From 1998 to 2011, the effects of environmental conditions on the spatial and temporal trends of sardine and sardinella catch rates in the Mauritanian waters were investigated using generalized additive models. Two models were used: a global model and an oceanographic model. The global models explained more of the variability in catch rates (60.4% for sardine and 40% for sardinella) than the oceanographic models (42% for sardine and 32.4% for sardinella). Both species showed clear and inverse seasonal variations in abundances corresponding to their main spawning activities and the hydrologic seasons off the Mauritanian waters. Sardine prefer colder waters and seem to occupy the ‘gap’ in the northern part of the Mauritanian waters as soon as sardinella has left the area because of to lower water temperatures. Unlike sardinella, sardine showed a gradual southward extension between 2002 and 2009. The oceanographic model revealed that a high proportion of catch variability for the two species could be explained by environmental variables. The main environmental parameters explaining the variability are sea surface temperature (SST) and the upwelling index for sardinella, and the chlorophyll‐a (Chl‐a) concentration, the upwelling index and SST for sardine. The sardinella spatio‐temporal variations off Mauritania seem to be more controlled by thermal than productivity gradients, probably linked to the species physiological constraints (thermal tolerance) whereas sardine seems to be more controlled by an ‘optimal upwelling and temperature’ windows. The results presented herein may be useful for understanding the movement of these species along the Mauritanian coast and hence their management under a changing climate.     

Population ecological parameters and biomass of anchovy kilka <Emphasis Type="Italic">Clupeonella engrauliformis</Emphasis> in the Caspian Sea

ABSTRACT:   Through most of the last century, three endemic kilka species supported major commercial species in the Caspian Sea. It is clear that catches and abundance of all species have changed, but catch and sampling data are limited and stock assessments are inadequate. Recent changes in the Caspian Sea ecosystem have occurred as a consequence of climatic environmental change (sea level change) and ecologic change caused by the invasive ctenophore Mnemiopsis leidyi . This paper examines the effects of these changes on the population biology and biomass of anchovy kilka Clupeonella engrauliformis in Iranian waters of the Caspian Sea from 1995 to 2004. For most years during this 10-year period, we estimated the age structure of catch, length–weight relationship, von Bertalanffy growth parameters, condition factors, sex ratios, maturity stages determined from ovarian analysis, natural and fishing mortality, age at first capture, and spawning biomass. The instantaneous coefficient of natural mortality was estimated as 0.473/year and the instantaneous coefficient of fishing mortality varied during the 10-year period between 0.541 and 2.690/year. Biomass of anchovy kilka declined from about 186 000 t in 1996 to less than 12 000 t in 2004. Recent high fishing rates were not sustainable after the introduction of Mnemiopsis , so overfishing is part of the explanation for the collapse of anchovy kilka in the Caspian Sea.     

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.     

Unveiling the influence of the environment on the migration pattern of the Atlantic pomfret (Brama brama) in North‐eastern Atlantic waters

Hydroclimatic variability is one of the main factors that drives inter‐annual changes in fish migration patterns. This study analyses the relationship between climate‐oceanographic factors and migration of the Atlantic pomfret (Brama brama) in NE Atlantic waters. Geo‐referenced catch data from logbooks of longliners operating in European Atlantic waters from 2002 to 2013 were linked to environmental indices at different temporal and spatial scales. Our results point to a strong influence of temperature at 200 m depth as the key factor along with the upwelling in the Galician (NW Iberian) waters. However, sea surface temperature (SST) indirectly affects the geographical display of Atlantic pomfret migration, and large migrations are observed in scenarios of high SSTs in the migratory area (c. above 14.7°C). Migrations are constrained during years when temperatures are below this threshold. A longer time‐series of annual landings (1950–2013) supports this evidence and highlights the significant influence of temperature at 200‐m depth along with the landings of the previous year. Length frequency distributions suggest an increase in size between consecutive seasons supporting the hypothesis that migration is a feeding strategy and a return to tropical waters of origin for spawning. Our study shows that the temperature of intermediate waters is a key variable in determining the northward migration of the Atlantic pomfret whereas density‐dependence and surface climatic conditions trigger secondary effects on the migration pattern of this species.     

Interannual variation in pelagic juvenile rockfish (Sebastes spp.) abundance – going with the flow

We report results from 28 yr of a midwater trawl survey of pelagic juvenile rockfish (Sebastes spp.) conducted off the central California coast. The fishery‐independent survey is designed to provide pre‐recruit indices of abundance for use in groundfish stock assessments. Standardized catch rate time series for 10 species were developed from delta‐generalized linear models that include main effects for year, station, and calendar date. Results show that interannual fluctuations of all 10 species are strongly coherent but highly variable, demonstrating both high‐ and low‐frequency components. A similarly coherent result is observed in the size composition of fish, with large fish associated with elevated catch rates. In contrast, spatial and seasonal patterns of abundance show greater species‐specific differences. A comparison of the shared common trend in pelagic juvenile rockfish abundance, derived from principal components analysis, with recruitments from five rockfish stock assessments shows that the time series are significantly correlated. An examination of oceanographic factors associated with year‐to‐year variability indicates that a signature of upwelled water at the time of the survey is only weakly related to abundance. Likewise, basin‐scale indices (the Multivariate El Niño‐Southern Oscillation Index, the Pacific Decadal Oscillation, the North Pacific Gyre Oscillation, and the Northern Oscillation Index) are poorly correlated with abundance. In contrast, sea level anomalies in the months preceding the survey are well correlated with reproductive success. In particular, equatorward anomalies in the alongshore flow field following the spawning season are associated with elevated survival and poleward anomalies with poor survival.     

Effect of wind stress on the catch of Japanese anchovy Engraulis japonicus off northwestern Kyushu, Japan

Effect of wind stress on the annual catch of Japanese anchovy Engraulis japonicus off northwestern Kyushu for the period between 1963 and 2009 was investigated. Regime shift analysis detected several step changes in catch and environmental variables. Since the mid-1980s, the anchovy catch in the coastal fishery zones has declined, while the catch in the offshore zone has increased. The decline of catch in the coastal zones showed a significant correlation with the long-term variations in prevailing north-northeastward wind stress over the Goto-Nada Sea during spring spawning season. The results indicated that weakened north-northeastward winds caused the recent low recruitment of anchovy through low levels of wind-induced eggs and larval transport from the offshore spawning ground to the coastal nursery areas, resulting in the potential shift of nursery area to the northwestern offshore region. Thus, as well as the growth-favorable ambient temperature, transport process would play a key role on long-term fluctuations in anchovy abundance in these coastal seas.     

Environmental conditions conducive to anchovy (Engraulis encrasicolus) spawning in the Adriatic Sea

The goal of this paper is to investigate the relationship between environmental variables and the occurrence of anchovy (Engraulis encrasicolus) spawning over a year, in order to enable the timing and strength of high spawning activity to be predicted. Biological sampling of anchovy was conducted on the fishing grounds of the eastern Adriatic (the northernmost semi‐enclosed basin in the Mediterranean) on a monthly basis from January 1999 to December 2010, while data on environmental parameters were taken from in situ measurements at the most representative and best surveyed eastern Adriatic hydrographic station and from I‐COADS monthly surface fields. Anchovy spawning seasonality was defined by monthly changes of the maturity stages and the gonadosomatic index. Fluctuations of the gonadosomatic index revealed that spawning begins in March and lasts until September, with reproductive activity peaking from April to July. A significant correlation was found between the gonadosomatic index and upper layer salinity in the two preceding months, while the lagged wind‐mixing index and surface temperature are barely correlated to the gonadosomatic index. An enhanced input of nutrient‐rich freshwater of river origin, which reduces upper layer salinity and enhances primary production up to 2 months before anchovy spawning, seems to be correlated with the anchovy fertility, especially for a late spawning maximum (in July).     

Catch rate and at‐vessel mortality of circle hooks versus J‐hooks in pelagic longline fisheries: A global meta‐analysis

We conducted a meta‐analysis of literature reporting on the use of circle hooks and J‐hooks in pelagic longline fisheries. Our study included more data than previous meta‐analyses of the effects of hook type, due to both a larger number of relevant studies available in recent years and a more general modelling approach. Data from 42 empirical studies were analysed using a random effects model to compare the effects of circle hooks and J‐hooks on catch rate (43 species) and at‐vessel mortality (31 species) of target and bycatch species. Catch rates with circle hooks were greater for 11 species, including four tuna species, six shark species and one Istiophorid billfish. Catch rates on circle hooks were lower for seven species, including two Istiophorid billfishes and two species of sea turtle. At‐vessel mortality was significantly lower with circle hooks in 12 species, including three tuna species, three Istiophorid billfishes, swordfish (Xiphias gladius) and three shark species. No species had significantly greater at‐vessel mortality when captured with a circle hook rather than a J‐hook. While our general approach increased model variability compared to more detailed studies, results were consistent with trends identified in previous studies that compared the catch rates and at‐vessel mortality (between hook types) for a number of species. Our results suggest that circle hooks can be a promising tool to reduce mortality of some bycatch species in pelagic longline fisheries, although the effects depend on the species and the metric (catch rate or at‐vessel mortality), emphasizing the need for fishery‐specific data in conservation and management decisions.     

Stock assessment and management implications of anchovy kilka (Clupeonella engrauliformis) in Iranian waters of the Caspian Sea

Recent changes in sea level of the Caspian Sea and ecological impacts caused by the invasive ctenophore (Mnemiopsis leidyi) have altered the ecosystem. A consequence is the changes in the absolute and relative abundance of the commercially important anchovy kilka (Clupeonella engrauliformis) in Iranian waters. To adjust to this change more rigorous management of this fishery is required. This paper examines the maximum sustainable yield (MSY) and fishing intensity at MSY. The paper presents estimates of f_MSY, yield-per-recruit and spawning biomass-per-recruit under various harvest strategies of F_max, F_0.1 and F_40%. We propose a method for estimating acceptable biological catch (ABC) that accounts for large differences in the quality and quantity of information and available data. The MSY and f_MSY were estimated 44,652 mt (metric tons) and 18,609 vessel × nights (a unit of effort). The ABC was estimated at 2190 mt in 2004. In 2005, however, the catch of anchovy kilka was about 4300, over twice the estimated ABC. In 2008 (from January to October) the catch declined to 220 mt. The analyses indicate that overfishing, especially between 2005 and 2008, is the main reason of the collapse of anchovy kilka in the Caspian Sea.     

Relative influence of static and dynamic features on black‐footed albatross (Phoebastria nigripes) habitat use in central California Sanctuaries

Effective conservation of highly mobile species requires an understanding of the factors that influence their habitat use patterns, locally and within a large‐scale oceanographic context. We characterized the seasonal (chick‐rearing, post‐breeding) and inter‐annual (2004–2008) distribution and abundance of black‐footed albatross (Phoebastria nigripes; BFAL) along the central California continental shelf/slope using standardized vessel‐based surveys. We used a hypothesis‐based information‐theoretic approach to quantify the relative influence of environmental conditions on BFAL occurrence and abundance by assessing their association with: (i) local static bathymetric features, (ii) local and regional dynamic oceanographic processes, and (iii) seasonal and inter‐annual basin‐wide variability. While the presence/absence models yielded stronger results than the abundance models, both revealed that static and dynamic features influence BFAL habitat use. Specifically, occurrence was greatest near the shelf‐break, particularly in months with strong upwelling. High BFAL densities were associated with Rittenburg Bank, especially during the chick‐rearing season, periods of positive North Pacific Gyre Oscillation index and large northern monthly upwelling, evidenced by cool, salty waters in the study area. BFAL aggregation intensity was greatest onshore of the shelf‐break (200 m isobath). Behavioral observations reinforced the notion that transiting BFAL are widely dispersed near the shelf‐break and concentrate in large flocks of birds sitting on the water farther onshore. These results underscore the need to consider oceanographic processes at multiple spatial scales when interpreting changes in BFAL dispersion within marine sanctuaries, and highlight the feasibility of implementing bathymetrically defined protected areas targeting predictable BFAL aggregations within these larger management jurisdictions.     

A biophysical model to assess the trade‐off between larval recruitment and catch in southern Australia's largest prawn fishery

Data from stock assessment surveys, published research and climate sensors were linked to model the interaction between fishing, physical‐oceanographic processes and spatial patterns of larval settlement for western king prawn [Penaeus (Melicertus) latisulcatus]. This information was used to evaluate the trade‐off between larval recruitment and catch during fishing periods that demand high prices but coincide with spawning. Total rates of larval settlement were maximized when tidal currents and atmospheric physical‐forcing components were coupled with simulations of larval swimming behaviour under average gulf temperatures. Average gulf temperatures sustained longer larval durations and increased larval settlement rates by over 12% compared with warmer gulf conditions simulated under a scenario of global warming. Reproductive data coupled with outputs from the biophysical model identified consistent inter‐annual patterns in the areas contributing to larval settlement success. Areas located in the north‐east, and central‐west of the fishery, consistently contributed to over 40% of all larvae reaching a settlement in each year. Harvest sensitivity analyses indicated that changes in the spatial patterns of pre‐Christmas fishing could lead to improvements in overall rates of the larval settlement while maintaining or improving the levels of catch. Future studies to refine the model inputs relating to physical processes, larval behaviour and mortality rates for P. latisulcatus coupled with surveys of juvenile prawn abundance to ground truth the modelled predictions, would allow stock recruitment relationships to be more closely examined and inform adaptive management of the fishery in the future.     

Brandt's cormorant diet (1994–2012) indicates the importance of fall ocean conditions for northern anchovy in central California

Effective ecosystem‐based management requires a comprehensive understanding of the functional links in the system. In many marine systems, forage species constitute a critical link between primary production and upper trophic level marine predators. As top predators, seabirds can be indicators of the forage species they consume and the ocean processes that influence these populations. We analyzed the diet and breeding success for the years 1994, 2003, 2005, and 2007–2012 of the Brandt's cormorant (Phalacrocorax penicillatus), a piscivorous diving seabird, breeding in central California, to evaluate the extent to which cormorant diet composition relates to prey availability, and how diet composition relates to breeding success and ocean conditions. Cormorant diet was primarily composed of young‐of‐the‐year (YOY) northern anchovy (Engraulis mordax), YOY rockfish (Sebastes spp.), and several species of small flatfish (order Pleuronectiformes). YOY rockfish consumption was positively related to their abundance as measured in a late spring pelagic midwater trawl survey. Northern anchovy appeared to be the most important prey as its consumption was positively related to cormorant breeding success. More northern anchovy were consumed in years where warm‐water conditions prevailed in the fall season before cormorant breeding. Thus, warm ocean conditions in the fall appear to be an important contributing factor in producing a strong year‐class of northern anchovy in central California and consequently a strong‐year class of Brandt's cormorant on the Farallon Islands.     

Disentangling density‐dependent effects on egg production and survival from egg to recruitment in fish

Understanding of density‐dependent effects is key to achieving sustainable management of self‐regulating biological resources such as fish stocks. Traditionally, density‐dependent effects on population abundance in fish have been considered to occur from hatching to recruitment, based on the paradigm of proportionality between spawning stock biomass and total egg production. Here, we demonstrate how the existence of intraspecific and interspecific density dependence in egg production changes the current understanding of density‐dependent processes in the life history of fish, by disentangling density‐dependent effects on egg production and survival from egg to recruitment, using sardine (Sardinops melanostictus, Clupeidae) and anchovy (Engraulis japonicus, Engraulidae) as model species. For sardine, strong intraspecific density‐dependent effects occurred in egg production, but no density‐dependent effects occurred or if any they were weak enough to be masked by environmental factors from hatching to recruitment. In contrast, for anchovy, interspecific density‐dependent effects occurred in egg production. In the survival after hatching, anchovy experienced stronger intraspecific density‐dependent effects than currently recognized. This analysis could overturn the current understanding of density‐dependent effects in the life history, highlighting contrasts between the effects on individual quality and population abundance and between the model species. We propose to reconsider the basis of fisheries management and recruitment studies based on the revised understanding of density‐dependent effects in the life history of the respective species.     

Biological investigations of anchovy and some selected fish species caught during the R／V “Bei Dou”surveys 1984—1999

IntroductionProductivity of an ecosystem may be measured in terms of structure,abundance and growth of keyspecies.Stock structure,measured as age or size composition in a given stock may indicate whether the stockis in a state of growing,stable or declining.Individual growth may be measured in terms of annual growth ofindividual age groups and maturity of key species will similarly reflect the potential of the species to copewith the influence of a changing environment of predators and human i…     

鳀鱼及其他重要经济鱼类生物学特征的研究

The paper deals with growth of Japanese anchovy (Engraulis japonicus) and variations in length by area and maturity of anchovy and some selected fish species obtained during surveys with R/V “Bei Dou” in the Yellow Sea and East China Sea,in 1984-1999.Demersal trawl catch rates for some selected fish species from the same surveys are also included.No density-dependent growth in length or weight of anchovy was found.Reduced length at maturity of small yellow croaker was apparent from 1986 to 1994.     

A generalized linear mixed model analysis of a multi-vessel fishery resource survey

A generalized linear mixed model (GLMM) that treats year and spatial cell as fixed effects while treating vessel as a random effect is used to examine fishing power among chartered industry-based vessels and a research trawler, the FRV Miller Freeman, for bottom trawl surveys on the upper continental slope of U.S. West coast. A Bernoulli distribution is used to model the probability of a non-zero haul and the gamma distribution to model the non-zero catch rates of four groundfish species. The use of vessel as a random effect allows the data for the various vessels to be combined and a single continuous time-series of biomass indices to be developed for stock assessment purposes. The GLMMs fit the data reasonably well. Among the different models examined, the GLMM incorporating a random vessel × year effect had the smallest ΔAIC and was thus chosen as the best model. Also, estimated random effects coefficients associated with the industry-based vessels and the FRV Miller Freeman for each year suggests that these vessels can be assumed to be from a common random effects distribution. These results suggest that combining data from the chartered industry-based vessels and from the research trawler may be appropriate to develop indices of abundance for stock assessment purposes. Finally, an evaluation of variances associated with abundance indices from the different models indicate that analyzing these data as a fixed effect GLM may underestimate the level of variability due to ignoring the grouped nature of tows within vessels. As such, use of a mixed model approach with vessel as a random effect is a reasonable approach to developing abundance indices and their variances.