Environmental and spatial effects on the distribution of blue marlin (Makaira nigricans) as inferred from data for longline fisheries in the Pacific Ocean

Blue marlin is distributed throughout tropical and temperate waters in the Pacific Ocean. However, the preference of this species for particular habitats may impact its vulnerability to being caught. The relationship between spatio‐temporal patterns of blue marlin abundance and environmental factors is examined using generalized additive models fitted to catch and effort data from longline fisheries. The presence of blue marlin, and the catch rate given presence, are modeled separately. Latitude, longitude, and sea‐surface temperature explain the greatest proportion of the deviance. Spatial distributions of relative density of blue marlin, based on combining the probability of presence and relative density given presence, indicate that there is seasonal variation in the distribution of blue marlin, and that the highest densities occur in the tropics. Seasonal patterns in the relative density of blue marlin appear to be related to shifts in SST. The distribution and relative abundance of blue marlin are sufficiently heterogeneous in space and time that the results of analyses of catch and effort data to identify ‘hotspots’ could be used as the basis for time‐area management to reduce the amount of blue marlin bycaught in longline fisheries.     

Importance of the Straits of Florida spawning ground to Atlantic sailfish (Istiophorus platypterus) and blue marlin (Makaira nigricans)

Much of the uncertainty in managing highly migratory pelagic species results from the scarcity of fisheries-independent data relevant to determining long-term trends in abundance, migratory movements, and the relative importance of different spawning grounds. To address these issues, we used an ichthyoplankton-based method to quantify the overall level of spawning of sailfish ( Istiophorus platypterus ) and blue marlin ( Makaira nigricans ) in the Straits of Florida (SF). We estimated that during the 2 years (2003–2004) of the study, 4.60 × 10¹¹ sailfish eggs and 4.49 × 10¹¹ blue marlin eggs were produced on an annual basis in this region. These egg production values, when combined with estimates of annual fecundity for each species and the most recent stock assessment estimate of total biomass, indicate that about 2.1% of Western Atlantic sailfish spawning and 1.6% of Atlantic-wide blue marlin spawning occurs in the SF. Additionally, pop-up satellite tags deployed on sailfish at the start of the spawning season revealed their short residency times in the SF, suggesting that a large (≈13%) transient portion of the sailfish population is responsible for the SF egg production. Overall, this study provides a critically needed fisheries-independent method of quantifying spatial and temporal trends in the abundance of highly migratory species. The application of this methodology in the SF indicated that above-average levels of sailfish and blue marlin spawning occur in this area and, possibly more importantly, that the SF is a migratory bottleneck for these species.     

Sex-structured population dynamics of blue marlin Makaira nigricans in the Pacific Ocean

The population dynamics of the blue marlin Makaira nigricans stock in the Pacific Ocean were estimated for 1971–2011 using a fully integrated length-based, age-, and sex-structured model. Fishery-specific catch, size composition, and catch-per-unit of effort were used in the modeling as likelihood components. Estimated dynamics were consistent with a stock that is fully exploited and stable over the last several years. No significant trends in recruitment were noted; however, female blue marlin were estimated to make up a majority of the catch, and historical exploitation has disproportionately changed the age structure of females relative to males. This result is due to differences in assumed life history and estimated selectivity. Changes to important life history parameters that are responsible for the productivity of the stock would potentially change the interpretation of current stock status.     

Spatial variability of bigeye tuna habitat in the Pacific Ocean: Hindcast from a refined ecological niche model

Habitat distribution is critically informative for stock assessment, since incorporating its variabilities can have important implications for the estimation of stock biomass or the relative abundance index. A refined ecological niche model with habitat characteristic parameterization was developed to reconstitute a 3‐D ecological map of bigeye tuna in the Pacific Ocean. We determined the boundaries and hierarchies of oceanographic features and hydrological conditions at horizontal and vertical scales to define the habitat preference of bigeye tuna associated with their feeding and physiological requirements. Ecogeographic projections underlined the depth‐ and region‐specific habitat distribution of bigeye tuna, with noticeable dynamic variations in the response to climate variability. Depths from 300 to 400 m represented layers of the most productive habitat, which was widespread through the equatorial Pacific Ocean and extended to the north‐central Pacific Ocean. The proportion of high‐quality habitat size in the north Pacific had a strictly regular intra‐annual cycle with peaks during the winter. Climate variability appeared to disturb the balance of the regular fluctuations in habitat size in the equatorial Pacific. Habitat hotspots during an El Niño period were characterized by their expansion to the north of the Hawaiian islands, shrinkage in the west for the hotspot band north of the Equator, and an eastern shift for the band south of the Equator. This variability may be the consequence of the incorporated fluctuations of the oxygen minimum zones (OMZ), current systems, and stratification in the open ocean.     

Habitat preferences of striped marlin (Kajikia audax) in the eastern Pacific Ocean

Striped marlin (Kajikia audax) is an epipelagic species distributed in tropical and temperate waters of the Pacific Ocean. In the central and eastern Pacific Ocean, it is captured principally in commercial longline fisheries, and in small artisanal fisheries, however, it is also taken throughout its range in this region as an incidental catch of the tuna purse‐seine fishery. Previous studies suggest that overexploitation and climate change may reduce abundance and cause changes in spatial distributions of marine species. The main objective of this study was to describe the habitat preferences of striped marlin and the changes in its distribution in response to environmental factors. Habitat modeling was conducted using a maximum entropy model. Operational level data for 2003–2014, collected by scientific observers aboard large purse seine vessels, were compiled by the Inter‐American Tropical Tuna Commission and were matched with detailed (4 km) oceanographic data from satellites and general circulation models. Results showed that the spatial distribution of habitat was dynamic, with seasonal shifts between coastal (winter) and oceanic (summer) waters. We found that the preferred habitat is mainly in coastal waters with warm sea surface temperatures and a high chlorophyll‐a concentration.     

Reproductive biology of blue marlin <Emphasis Type="Italic">Makaira nigricans</Emphasis> around Yonaguni Island,southwestern Japan

We studied the spawning seasonality and gonadal development of blue marlin Makaira nigricans using specimens captured around Yonaguni Island in southwestern Japan between February 2003 and February 2006. The mean (±SD) lower jaw–fork length of females (234 ± 24 cm) was greater than that of males (191 ± 12 cm). The smallest mature female and male were 183 and 160 cm, respectively. Most of the 717 females had immature ovaries. However, in March and from May to September, the ovaries of 26 females contained oocytes with yolk globules, hydrated oocytes, or postovulatory follicles. Most males had testes with a large amount of spermatozoa throughout the year. The occurrence of mature blue marlin at Yonaguni Island suggests that spawning occurs here. The mean condition factors (fatness of the fish) of both sexes decreased from March to June or July, presumably as they expended energy to reproduce. We discuss our results in the context of migration theory for blue marlin in the western North Pacific Ocean.     

The population dynamics of the longnose skate,Raja rhina,in the northeast Pacific Ocean

The longnose skate, Raja rhina, is the most commonly landed skate species in the northeast Pacific Ocean. It also dominates survey catches in the area. Existing knowledge on skate biology and fisheries suggests that large skate species with late sexual maturation, like the longnose skate, are extremely vulnerable to overfishing, and if overfished they are slow to recover. Historically, in the US Pacific Coast skates have not supported directed fisheries, but have been taken as bycatch in other commercially important fisheries. For the past 13 years, landed catches of skates in waters off Washington, Oregon and California have significantly increased. Using the Stock Synthesis 2 modeling framework, a population model for the longnose skate was developed to assess the current state of the stock and reconstruct its past dynamics. Model results indicate that the longnose skate spawning biomass has slowly declined since 1915; the current stock biomass is estimated as 66% of its unexploited level. Uncertainty in regards to fishery-dependent data and female maturity parameters was explored through sensitivity analyses. The model results indicate that it is important to conduct species-specific identification in the fishery and monitor discard of longnose skate to improve the accuracy of fishery catch data and make assessment conclusions more robust.     

How consistent is the advice from stock assessments? Empirical estimates of inter-assessment bias and uncertainty for marine fish and invertebrate stocks

Fishery management frequently involves precautionary buffering for scientific uncertainty. For example, a precautionary buffer that scales with scientific uncertainty is used to calculate the acceptable biological catch downward from the overfishing limit in the US federal fishery management system. However, there is little empirical guidance to suggest how large buffers for scientific uncertainty should be. One important component of uncertainty is variation among different assessments of the same stock in estimates of management-relevant quantities. We analysed commercially exploited marine fish and invertebrate stocks around the world and developed Bayesian hierarchical models to quantify inter-assessment variation in terminal year biomass and fishing mortality estimates, reference points, relative biomass and fishing mortality estimates, and overfishing limits. There was little evidence of inter-assessment bias; stock assessment estimates in the terminal year of the assessment were not consistently higher or lower than estimates of the same quantities in future years. However, there was a tendency for extreme values from the terminal year to be pulled closer to the mean in future years. Inter-assessment variation in all estimates differed across regions, and a longer inter-assessment interval generally resulted in greater variation. Inter-assessment uncertainty was greatest for estimates of the overfishing limit, with coefficients of variation ranging from 17% in Europe (non-EU) to 107% for Pacific Ocean pelagic stocks. Because inter-assessment variation is only one component of scientific uncertainty, we suggest that these uncertainty estimates may provide a basis for determining the minimum size of precautionary buffers.     

Multi‐decadal variations in spawning ground use in Northeast Arctic haddock (Melanogrammus aeglefinus)

Despite the importance both in an ecological and management context, much uncertainty remains about the underlying factors controlling spawning ground use in marine fish. Here, we investigate how spawning ground use of Northeast Arctic (NEA) haddock (Melanogrammus aeglefinus) has changed over time. By combining data from a Soviet‐Russian egg survey conducted over a 35 year period (1959–1993) with a biophysical drift model, we estimate annual median spawning latitudes of NEA haddock. Furthermore, we test for statistical association between the median latitude of spawning and covariates that are candidates for shaping the spawning distribution of haddock. The results indicate no association with temperature, as measured at the Kola transect, or demographic structure, as measured by the mean weight of spawners. However, we find a positive and significant association between spawning latitude and spawning stock biomass, indicating that density‐dependence may be more important in shaping spawning location than other factors such as temperature and demographic structure. A potential mechanism for the reported effect of spawning stock biomass may act through the distribution at the feeding grounds, which is reported to expand north‐ and eastwards at high population density.     

Abundance estimation of Northeast Atlantic mackerel based on tag recapture data—A useful tool for stock assessment?

In the present study we utilize tag recapture data to estimate year class abundance and spawning stock biomass of mackerel (Scomber scombrus L.) in the Northeast Atlantic for the period 1986–2008. On average 20,000 jigged mackerel have been tagged annually with internal steel tags in the spawning area west of Ireland and the British Isles, and the tags have been recaptured in commercial catches screened through metal detectors. The spawning stock biomass estimates derived from two different tag-based models were highly variable but were on average 2 and 2.3 times higher than the ICES official estimate. The official estimate is considered uncertain and most likely an underestimate of the actual biomass, due to unregistered mortality in the fisheries and lack of fishery-independent, age-disaggregated data. Hence, tag-based estimates could potentially improve the current assessment if included in the ICES stock assessment on a regular basis. These estimates also involve some uncertainty that needs consideration, especially related to variable tagging mortality, detector efficiency and migrations of the stock.     

Influence of temperature and oxygen on the distribution of blue marlin (Makaira nigricans) in the Central Pacific

The blue marlin (Makaira nigricans) is a highly migratory pelagic predator of tropical and subtropical seas. Information on the habitat use of marine species is fundamental to understanding their ecology and population dynamics and is needed to inform responsible management strategies. Using a long‐term satellite tagging data set from The International Game Fish Association Great Marlin Race, we examined habitat use and how oxygen and temperature influence the horizontal and vertical distributions of blue marlin in the Central Pacific. Blue marlin primarily occurred in warm waters (26–30°C) and exhibited a diel bimodal depth distribution across the 5‐year data record (2009–2013), with fish spending the majority of their time near the surface at night and at deeper depths during the day (25–100 m). The depth distribution of blue marlin was limited in areas where low oxygen and/or temperature conditions occur closer to the surface, with the extent of habitat compression being greatest when both oxygen and temperature were limiting. The migrations of blue marlin appeared restricted during the 2010 La Niña, when increased equatorial upwelling resulted in an extension of the cold, low oxygen waters of the cold tongue into the Central Pacific, creating a barrier to the trans‐equatorial migrations that occurred during all other tagging years. If the frequency and intensity of La Niña events increases and the oxygen minimum layer continues to expand as has been predicted under certain climate change scenarios, the migratory behavior and habitat availability of blue marlin may be impacted.     

Basin‐scale reproductive segregation of Pacific halibut (Hippoglossus stenolepis)

Pacific halibut Hippoglossus stenolepis (Schmidt) is presently considered to consist of a single spawning population extending from California through the Bering Sea. However, this satellite tagging investigation suggests that geographic landforms and discontinuities in the continental shelf appear to limit the interchange of mature Pacific halibut among large marine ecosystems and delineate the boundaries of potential spawning components in the Gulf of Alaska and Bering Sea, with smaller components along the Aleutian Islands. The geographic segregation of these spawning components may be reinforced by regional behavioural adaptations and different temperature regimes in each area. These results suggest that the Pacific halibut population may be segregated into somewhat discrete spawning units among which less mixing is likely than that which occurs within them. As such, future stock assessment metrics may be most effective in preserving population function if spawning ecology is treated as a basin‐scale process.     

Density dependence in total egg production per spawner for marine fish

A paradigm of fisheries science holds that spawning stock biomass (SSB) is directly proportional to total egg production (TEP) of fish stocks. This “SSB–TEP proportionality” paradigm has been a basic premise underlying the spawner–recruitment models for fisheries management and numerous studies on recruitment mechanisms of fish. Studies on maternal effects on reproductive potential of a stock have progressed during the last few decades, leading to doubt concerning the paradigm. Nonetheless, a direct test of the paradigm at multidecadal scales has been difficult because of data limitations in the stock assessment systems worldwide. Here, we tested the paradigm for marine fish based on a novel combination of two independent 38‐year time series: fishery‐dependent stock assessment data and fishery‐independent egg survey data. Through this approach, we show that the SSB–TEP proportionality is distorted by density dependence in total egg production per spawner individual (TEPPS) or spawner unit weight (TEPPSW) at a multidecadal scale. The TEPPS/TEPPSW exponentially declined with biomass and thus was density‐dependent for Japanese sardine, a small pelagic species exhibiting a high level of population fluctuation, in the western North Pacific. By contrast, the TEPPS/TEPPSW was sardine‐density‐dependent for Japanese anchovy, another small pelagic species exhibiting a moderate level of population fluctuation well‐known for being out of phase with sardine. Our analysis revealed intraspecific (sardine) and interspecific (anchovy) density dependence in TEPPS/TEPPSW, which was previously unaccounted for in spawner–recruitment relationships. Such density‐dependent effects at the time of spawning should be considered in fisheries management and studies on recruitment mechanisms.     

Temporal variation of seasonality of egg production and the spawning biomass of Pacific anchovy, Engraulis japonicus, in the southern waters of Korea in 1983–1994

Embryonic mortality, egg production and the spawning stock biomass of Pacific anchovy, Engraulis japonicus , off Southern Korea during 1983–1994, and their biological response to oceanographic features in spring and summer, were analysed. The instantaneous mortality rate (IMR) of embryonic stages decreased in spring and increased in summer, with a range of 0.33–1.23 day^–1 in spring and 0.78–1.69 day^–1 in summer. Egg production in summer was three times that during spring and production was low in the late 1980s. Mean lengths of yolk-sac larvae and adult females were greater in spring than in summer, whereas spawning fraction and spawning stock ratio (spawning biomass:adult biomass) were lower in spring than summer. Estimated mean spawning stock biomass ranged from 141 × 10³ to 380 × 10³ MT in spring and from 221 × 10³ to 557 × 10³ MT in summer. Statistically, the seasonal and long-term trends of embryonic mortality, egg production and spawning stock biomass of Pacific anchovy can be explained largely by spring warming, summer cooling and by less abundant zooplankton in the late 1980s.     

Migrations and movements of Atlantic tarpon revealed by two decades of satellite tagging

Understanding large‐scale migratory behaviours, local movement patterns and population connectivity are critical to determining the natural processes and anthropogenic stressors that influence population dynamics and for developing effective conservation plans. Atlantic tarpon occur over a broad geographic range in the Atlantic Ocean where they support valuable subsistence, commercial and recreational fisheries. From 2001 through 2018, we deployed 292 satellite telemetry tags on Atlantic tarpon in coastal waters off three continents to document: (a) seasonal migrations and regional population connectivity; (b) freshwater and estuarine habitat utilization; (c) spawning locations; and (d) shark predation across the south‐eastern United States, Gulf of Mexico and northern Caribbean Sea. These results showed that some mature tarpon make long seasonal migrations over thousands of kilometres crossing state and national jurisdictional borders. Others showed more local movements and habitat use. The tag data also revealed potential spawning locations consistent with those inferred in other studies from observations of early life stage tarpon leptocephalus larvae. Our analyses indicated that shark predation mortality on released tarpon is higher than previously estimated, especially at ocean passes, river mouths and inlets to bays. To date, there has been no formal stock assessment of Atlantic tarpon, and regional fishery management plans do not exist. Our findings will provide critical input to these important efforts and assist the multinational community in the development of a stock‐wide management information system to support informed decision‐making for sustaining Atlantic tarpon fisheries.     

Comparison of biomass estimates from multiple stratification approaches in a swept area method for Pacific saury Cololabis saira in the western North Pacific

Fishery-independent surveys using sea surface trawl nets for Pacific saury Cololabis saira in the western North Pacific since 2003 have enabled the investigation of their annual distribution patterns and total biomass during June and July, prior to the main fishing season in Japan. We compared biomass estimates and their associated variances derived based on five different post-stratification approaches in a swept area method, and then observed that these approaches have little effect on biomass estimates and their precision, owing to well-organized survey designs and homogeneously allocated sampling stations. We were able to utilize decreasing biomass estimates for 15 years as an essential abundance index in the ongoing stock assessment. Notably, examination based on stratification along with longitudinal survey lines indicated that the estimated biomass had decreased in the western survey area, resulting in an eastward shift in the gravity center of Pacific saury distribution after 2010. We recommend biomass estimation in an east–west direction based on longitudinal stratification as an effective measure to develop population dynamics models which reflect westward migration into the fishing grounds around Japanese coastal waters, and to forecast the expected catch during the subsequent fishing period.

     

Incorporating an environmental stock–recruitment relationship in the assessment of Pacific cod (Gadus macrocephalus)

The robustness of a previously described environmentally mediated stock–recruitment relationship for Pacific cod in Hecate Strait, BC, Canada was tested with 10 yr of additional data. The original analysis tested several alternative hypotheses and concluded that water transport through Hecate Strait, as indicated by sea level height, coupled with cod spawning biomass formed the best model. The present analysis indicates the relationship held through the 1990s. The implications of variation in sea level on stock production were investigated with a delay‐difference stock production model that included an environmentally mediated stock–recruitment relationship. The model predicted that the maximum fishery yield would vary between 1750 and 3670 t yr^?1 over the observed range of sea level height, and the estimated unsustainable fishing mortality during periods of low productivity would be only 0.5 times that in periods of high productivity.     

Spatial segregation of striped marlin (Kajikia audax) by size in the eastern Pacific Ocean

Striped marlin (Kajikia audax) is an epipelagic fish distributed in oceanic and coastal waters of the Pacific Ocean. This species is usually found in warm and coastal waters with high primary productivity. The main goal of this study was to describe the spatial segregation of striped marlin by average Eye‐Fork length (EFL) in the eastern Pacific Ocean (EPO) and its relationship with environmental variables using EFL data obtained from tuna purse‐seining and Generalized Additive Models (GAMs). The model suggested that larger individuals of striped marlin were more likely to be found in waters with high Chlorophyll‐a concentration (>2 mg/m³) and with temperatures lower than 25°C, within a region known as the “cold tongue” and the Humboldt current system, while smaller individuals were more likely to be found in warmer and low productive areas within a region known as the “warm pool of the EPO.” We observed that set type caused a large variation on average EFL of striped marlin; larger fish were captured in sets associated with floating objects (natural and manmade), while smaller fish were captured in sets associated with dolphins. Despite this, our findings suggest that striped marlin has a latitudinal gradient in average EFL; larger individuals occurred predominantly south of 10°N, while smaller ones occurred predominantly in coastal waters between 10°N and 20°N, thus demonstrating a spatial segregation of the species affected by its maturity stage.     

Dependence of recruitment on parent stock of the spiny lobster,Panulirus argus,in Florida

Despite management practices to achieve sustainability, commercial landings for Florida spiny lobster (Panulirus argus) have experienced a drastic decline (57%) since 2000. This is cause for concern not only for economic reasons, but for issues of sustainability. An annual index of P. argus post‐larval (puerulus) abundance, estimated with a generalized linear model with significant mean sea‐level effects, shows a 36% decrease in annual puerulus supply since 1988. In addition, local Florida spawning stock biomass estimated from an age‐structured sequential population analysis decreased 57% since 1988. Puerulus abundance follows a highly correlated (R = 0.76) trend with a 12‐month delayed spawning stock abundance, which supports the contention that the Florida spawning population is a significant contributing factor to post‐larval recruitment in Florida. Residuals about the puerulus on spawning stock abundance function follow closely an interannual North Atlantic Oscillation Index signal. This residual effect is thought of as a secondary regional population effect on Florida puerulus recruitment. The Florida spiny lobster stock is exploited with no fishing mortality controls due to the Pan Caribbean recruitment concept adopted in Florida spiny lobster management. Therefore, the potential of recruitment overfishing exists if fishing mortality controls to protect local spawning stock abundance, such as catch quotas, are not introduced.     

Distribution of albacore (Thunnus alalunga) in the Indian Ocean and its relation to environmental factors

The distribution pattern of albacore, Thunnus alalunga, in the Indian Ocean was analyzed based on catch data from the Taiwanese tuna longline fishery during the period 1979–85. The Taiwanese tuna fishery began operating in the Indian Ocean in 1967. We used a geographic information system to compile a fishery and environmental database and statistically explored the catch per unit effort (CPUE) distribution of albacore. Our results indicated that immature albacore were mainly distributed in areas south of 30°S although some displayed a north–south seasonal migration. Mature albacore, which were mainly concentrated between 10°S and 25°S, also showed a north–south migration. Within 10°S and 30°S, the separation of mature, spawning, and immature albacore life history stages roughly coincided with the boundaries of the three oceanic current systems in the Indian Ocean. The optimal environmental variables for CPUE prediction by stepwise discriminant analysis differed among life history stages. For immature albacore, the sea surface variables sea surface temperature (SST), chlorophyll concentration and surface salinity were significant. For mature albacore, SST was significant, while for spawning albacore, the sub‐surface variables temperature at 100 m and oxygen at 200 m were significant. Spawning albacore evidently prefer deep oceanographic conditions. Our results on the oceanographic conditions preferred by different developmental stages of albacore in the Indian Ocean were compatible with previous studies found in the Pacific Ocean.