The rostral micro-tooth morphology of blue marlin, Makaira nigricans.

Billfish rostra potentially have several functions; however, their role in feeding is unequivocal in some species. Recent work linked morphological variation in rostral micro-teeth to differences in feeding behavior in two billfish species, the striped marlin (Kajikia audax) and the sailfish (Istiophorus platypterus). Here, we present the rostral micro-tooth morphology for a third billfish species, the blue marlin (Makaira nigricans), for which the use of the rostrum in feeding behavior is still undocumented from systematic observations in the wild. We measured the micro-teeth on rostrum tips of blue marlin, striped marlin, and sailfish using a micro-computed tomography approach and compared the tooth morphology among the three species. This was done after an analysis of video-recorded hunting behavior of striped marlin and sailfish revealed that both species strike prey predominantly with the first third of the rostrum, which provided the justification to focus our analysis on the rostrum tips. In blue marlin, intact micro-teeth were longer compared to striped marlin but not to sailfish. Blue marlin had a higher fraction of broken teeth than both striped marlin and sailfish, and broken teeth were distributed more evenly on the rostrum. Micro-tooth regrowth was equally low in both marlin species but higher in sailfish. Based on the differences and similarities in the micro-tooth morphology between the billfish species, we discuss potential feeding-related rostrum use in blue marlin. We put forward the hypothesis that blue marlin might use their rostra in high-speed dashes as observed in striped marlin, rather than in the high-precision rostral strikes described for sailfish, possibly focusing on larger prey organisms.

Broadbill swordfish (Xiphias gladius) foraging and vertical movements in the north-west Atlantic.

The northern edge of Georges Bank is an important seasonal foraging habitat for swordfish (Xiphias gladius) in the North Atlantic, where aggregations support commercial pelagic longline and harpoon fisheries. Following a period of overfishing during the 1990s, the North Atlantic X. gladius stock underwent a period of recovery during the early 2000s and was considered rebuilt in 2009. We analysed stomach contents from X. gladius (n = 39) harvested by the Canadian harpoon fishery on Georges Bank in 2007 to characterize diet in this important foraging habitat. We used electronic tagging data from X. gladius (n = 6) on Georges Bank in 2005-2007 to assess vertical habitat preferences and associated prey composition within those zones. We also used stable isotope analysis (δ13 C and δ15 N) of X. gladius liver (n = 2) and common prey types (Paralepididae, Myctophidae, Merluccidae, Ommastrephidae) as a longer-term record of feeding. Stomach contents were co-dominated by Paralepididae [31.9% weight (W)] and Ommastrephidae (36.8%W) with secondary contributions from hake (Merluccidae, 6.5%W), Myctophidae (2.9%W) and Sebastidae (2.1%W). X. gladius displayed diel vertical migrations, descending to depths of 300-400 m during daytime followed by residence in surface waters at night. X. gladius liver δ15 N values were similar to or lower than values of primary stomach contents, likely due to bias of diet consumed in southerly waters with lower nitrogen isotope baselines prior to arrival on Georges Bank. Diet data are similar to results from historical studies from the late 1950s to the early 1980s. This apparent temporal stability to the underlying food web in this region may explain the high X. gladius site fidelity observed in electronic tagging studies and the consistent aggregation of these fish to this region.

Movement ecology of black marlin Istiompax indica in the Western Indian Ocean.

The black marlin Istiompax indica is an apex marine predator and is susceptible to overfishing. The movement ecology of the species remains poorly known, particularly within the Indian Ocean, which has hampered assessment of their conservation status and fisheries management requirements. Here, we used pop-up archival satellite tags to track I. indica movement and examine their dispersal. Forty-nine tags were deployed off Kenya during both the north-east (November-April) and south-west (August-September) monsoon seasons, providing locations from every month of the year. Individual I. indica were highly mobile and track distance correlated with the duration of tag attachment. Mean track duration was 38 days and mean track distance was >1800 km. Individuals dispersed in several directions: north-east into Somalian waters and up to northern Oman, east towards the Seychelles, and south into the Mozambique Channel. Their core habitat shifted seasonally and overlapped with areas of high productivity off Kenya, Somalia and Oman during the first half of the year. A second annual aggregation off the Kenyan coast, during August and September, did not coincide with high chlorophyll-a (chl-a) concentrations or thermal fronts, and the drivers of the species' presence and movement from this second aggregation was unclear. We tested their habitat preferences by comparing environmental conditions at track locations to the conditions at locations along simulated tracks based on the empirical data. Observed I. indica preferred cooler water with higher chl-a concentrations and stayed closer to the coast than simulated tracks. The rapid and extensive dispersal of I. indica from Kenya suggests that there is likely a single stock in the Western Indian Ocean, with individuals swimming between areas of high commercial catches off northern Somalia and Oman, and artisanal and recreational fisheries catches throughout East Africa and Mozambique.

Linking hunting weaponry to attack strategies in sailfish and striped marlin.

Linking morphological differences in foraging adaptations to prey choice and feeding strategies has provided major evolutionary insights across taxa. Here, we combine behavioural and morphological approaches to explore and compare the role of the rostrum (bill) and micro-teeth in the feeding behaviour of sailfish (Istiophorus platypterus) and striped marlin (Kajikia audax) when attacking schooling sardine prey. Behavioural results from high-speed videos showed that sailfish and striped marlin both regularly made rostrum contact with prey but displayed distinct strategies. Marlin used high-speed dashes, breaking schools apart, often contacting prey incidentally or tapping at isolated prey with their rostra; while sailfish used their rostra more frequently and tended to use a slower, less disruptive approach with more horizontal rostral slashes on cohesive prey schools. Capture success per attack was similar between species, but striped marlin had higher capture rates per minute. The rostra of both species are covered with micro-teeth, and micro-CT imaging showed that species did not differ in average micro-tooth length, but sailfish had a higher density of micro-teeth on the dorsal and ventral sides of their rostra and a higher amount of micro-teeth regrowth, suggesting a greater amount of rostrum use is associated with more investment in micro-teeth. Our analysis shows that the rostra of billfish are used in distinct ways and we discuss our results in the broader context of relationships between morphological and behavioural feeding adaptations across species.

Movement and habitat use of striped marlin Kajikia audax in the Western Indian Ocean.

Striped marlin Kajikia audax are globally Near Threatened and their stock in the Indian Ocean was last assessed as "overfished and subject to overfishing". Significant gaps in our understanding of their ecology remain, hampering the efforts of fisheries managers to ensure stock sustainability. There is a particular lack of fisheries-independent data. Here we present the results from the first large-scale satellite tracking study of K. audax in the Indian Ocean. We tagged 49 K. audax with pop-up archival satellite-linked tags off the Kenyan coast from 2015 to 2019. Individuals were highly mobile, covering horizontal distances of up to 9187 km over periods ranging up to 183 days, with a mean daily distance of ~48 km. Long-distance movements were recorded to the east and north of East Africa, with the most distant tracks extending north to the Arabian Sea and east to near the Maldives. None of the K. audax swam south of East Africa. Kernel utilization distributions of fish locations demonstrated their shifting seasonal activity hotspots. Over the sport-fishing season (and tagging period) in Kenya, from December to March, K. audax typically stayed off the East African coast. After March, the activity hotspot shifted north to a region close to the Horn of Africa and Socotra Island. Remotely sensed sea surface temperature and chlorophyll-a maps indicated that this seasonal movement could be driven by a shift in prey availability. Our results show the high mobility of K. audax in the Western Indian Ocean, and that individuals seasonally range between two major fishing areas.

Reproductive biology of female striped marlin Kajikia audax in the western Pacific Ocean.

Length and mass data for 1260 (536 females, 683 males, 41 sex unknown) striped marlin Kajikia audax were collected at the fish markets of Tungkang, Singkang and Nanfangao from July 2004 to September 2010. Of these samples, 534 gonads (236 females and 298 males) ranging from 95 to 206 cm in eye-to-fork length (LEF ) and 8 to 88 kg in round mass (MR ), were collected. Chi-square tests indicated sex ratios were homogeneous among months in 2004 and 2006-2008, but not in 2005, 2009 and 2010; and there were significant differences in sex ratio by size. The overall sex ratio (RS ) differed significantly from the expected 0·5. Kajikia audax are sexually dimorphic and the proportions of females increased with size between 140 and 210 cm LEF . Reproductive activity was assessed using a gonado-somatic index (IG ), external appearance of the gonads and histological examination and results indicated that the spawning season occurred from April to August with a peak in June to July. Based on histological observations and the distribution of oocyte diameters, K. audax are multiple spawners and their oocytes develop asynchronously. The estimated length-at-50% maturity (LEF50 ) was c. 181 cm (c. 4·8 years of age) for females. The proportion of reproductively active females in the spawning season with ovaries containing postovulatory follicles (0·27) indicated that they spawned every 3·7 days on average. The hydrated oocyte method estimated mean ± S.D. batch fecundity (FB ) to be 4·4 ± 2·02 million eggs; average relative fecundity was 53·6 ± 13·9 oocytes g-1 MR ; and the average annual fecundity was 181·3 ± 48·3 million eggs. The parameters estimated in this study are key information for stock assessments of K. audax in the north-western and central Pacific and will contribute to the conservation, management and sustainable yield of this species.

Genetic identification of istiophorid larvae from the Gulf of Mexico based on the analysis of mitochondrial DNA control region sequences.

Assigning relative importance of spawning and nursery habitats for threatened and endangered teleosts, such as those seen in the Gulf of Mexico (GoM), relies on the proper identification of the early life-history stages of the species of concern. Here, sequencing a portion of the mitochondrial DNA (mtDNA) control region (CR) I as barcodes is recommended to identify istiophorid (billfish) larvae in the Atlantic Ocean because of its high resolution and the intrinsic value of the levels of genetic variation that can be extracted from these data. The universality of the primers employed here demonstrates their utility for not only the positive identification of istiophorids in the GoM, but for any larval teleost occurring in areas recognized as larval hotspots worldwide.

Mercury in the Blue Marlin (Makaira nigricans) from the Southern Gulf of California: Tissue Distribution and Inter-Annual Variation (2005-2012).

With the aim of knowing annual variations of Hg concentrations in muscle and liver of blue marlin (Makaira nigricans) from the southern Gulf of California, fish were collected between 2005 and 2012 in three areas. In general, Hg levels were more elevated in liver than in muscle. Variations of Hg concentrations in muscle and liver among the studied years were not significant. Hg levels in muscle and liver increased significantly with length and weight of fish. In comparison to other studies, Hg levels in muscle and liver were lower. With respect to maximum permissible limits (1.0 µg g-1 wet weight) of Hg in Mexico, the average concentration (1.91 µg g-1 wet weight) in the edible portion (muscle) of blue marlin was over the legal limit; this issue is worth research efforts in relation to the rate of ingestion of this species and the co-occurrence of selenium in the edible portion.

Feeding in billfishes: inferring the role of the rostrum from a biomechanical standpoint.

Perhaps the most striking feature of billfishes is the extreme elongation of the premaxillary bones forming their rostra. Surprisingly, the exact role of this structure in feeding is still controversial. The goal of this study is to investigate the use of the rostrum from a functional, biomechanical and morphological standpoint to ultimately infer its possible role during feeding. Using beam theory, experimental and theoretical loading tests were performed on the rostra from two morphologically different billfish, the blue marlin (Makaira nigricans) and the swordfish (Xiphias gladius). Two loading regimes were applied (dorsoventral and lateral) to simulate possible striking behaviors. Histological samples and material properties of the rostra were obtained along their lengths to further characterize structure and mechanical performance. Intraspecific results show similar stress distributions for most regions of the rostra, suggesting that this structure may be designed to withstand continuous loadings with no particular region of stress concentration. Although material stiffness increased distally, flexural stiffness increased proximally owing to higher second moment of area. The blue marlin rostrum was stiffer and resisted considerably higher loads for both loading planes compared with that of the swordfish. However, when a continuous load along the rostrum was considered, simulating the rostrum swinging through the water, swordfish exhibited lower stress and drag during lateral loading. Our combined results suggest that the swordfish rostrum is suited for lateral swiping to incapacitate their prey, whereas the blue marlin rostrum is better suited to strike prey from a wider variety of directions.

Betanodavirus meningoencephalitis in an Atlantic blue marlin.

Viral nervous necrosis (viral encephalopathy and retinopathy) is caused by piscine nodavirus (Nodaviridae, Betanodavirus). Since 1986, this highly infectious virus has caused mass mortalities of up to 100% in farmed saltwater and freshwater fish around the world (with the exception of South America and Antarctica), affecting >60 species across 10 orders. The Atlantic blue marlin (Makaira nigricans Lacépède, 1802) is a top-level predator found throughout the tropical waters of the Atlantic and Indo-Pacific oceans. Despite their popularity as a sportfish, relatively little is known about the Atlantic blue marlin and other billfish. We describe here chronic betanodavirus infection in a juvenile Atlantic blue marlin, which is, to our knowledge, the first report of disease in M. nigricans.

Essential and non-essential elements in tuna and billfish around the world: Distribution patterns and influencing factors.

Tuna and billfish are widely distributed in oceans worldwide. Their survival is relied on a decent share of essential and non-essential elements. We conducted a comprehensive evaluation of essential and non-essential elements in livers of tuna and billfish collected from global oceans. The individual element consistently shown similar orders of magnitude in both tuna and billfish, with essential elements generally being 1-3 orders of magnitude higher than non-essential elements. Various physicochemical properties and behaviors contributed to four distinct clusters of these elements. Also, element distribution pattern indicated the presence of four sample groups based on regions and categories. Nine elements served as characteristic indicators. Among them, fish category was the most important influencing factor. Hg, Fe, Tl, Co, and Se were influenced by body size, trophic level, and feeding habits. Ni was influenced by sampling regions, while Mg, Mn and As were influenced by body size and local primary production.

Assessment of the Impact on 20 Pelagic Fish Species by the Taiwanese Small-Scale Longline Fishery in the Western North Pacific Using Ecological Risk Assessment.

Ecological risk assessment (ERA) has been applied on assessing the relative risk of bycatch species in recent years. ERA index is calculated by productivity of species and susceptibility of fisheries on fish species. In this study, a semi-quantitative method was used to evaluate the risks of exploitation for 20 pelagic fish species by the small-scale longline fisheries in the western North Pacific Ocean. The productivity was estimated based on the ranking (high, median, and low) of seven life history parameters. The susceptibility was calculated by the multiplication of the catchability, selectivity and post-capture mortality. The ERA results indicated the risks of sharks are higher than those of tunas and billfishes, except yellowfin tuna (Thunnus albacares). The shortfin mako shark (Isurus oxyrinchus) and dusky shark (Carcharhinus obscurus) have the highest risk. Other shark species, yellowfin tuna, and sailfish (Istiophorus platypterus) have medium risk. While the striped marlin (Kajikia audax), and albacore tuna (T. alalunga) have the lowest risk. Stock assessment and rigorous management measures such as catch quota and size limit are recommended for the species in high or medium ecological risk and a consistent monitoring management scheme is suggested for those in low ecological risk.

Status and challenges for sustainable billfish fisheries in the Western Indian Ocean.

Billfish species (families Istiophoridae and Xiphiidae) are caught in artisanal, recreational, and commercial fisheries throughout the Western Indian Ocean region. However, data and information on the interactions among these fisheries and the ecology of billfish in the WIO are not well understood. Using an in-depth analysis of peer-reviewed articles, grey literature, observation studies, and authors' insider knowledge, we summarize the current state of knowledge on billfish fisheries in 10 countries. To describe historical and current trends, we examined fisheries statistics from governmental and non-governmental agencies, sportfishing clubs' reports, diaries of sportfishing captains, and the catch and effort databases of the Indian Ocean Tuna Commission. We highlight two key points. First, billfish fisheries in the Western Indian Ocean are highly diverse, comprising two distinct segments-coastal and oceanic. However, data are poor for most countries with significant gaps in information especially for sport and artisanal fisheries. Second, the evidence assembled showed that billfish species have immense social, cultural, and economic value. Swordfish are targeted by both large-scale and semi-industrial fisheries, while other billfish species, particularly marlin, are highly sought after by sport fisheries in most countries. Our paper provides a comprehensive review of billfish fisheries and available information in the context of the WIO underscoring the need to strengthen data collection and reporting, citizen science, and collaborative sustainable development and management of billfish. Supplementary Information: The online version contains supplementary material available at 10.1007/s11160-022-09725-8.

Mercury, selenium, and stable carbon and nitrogen isotopes in the striped marlin Kajikia audax and blue marlin Makaira nigricans food web from the Gulf of California.

Mercury (Hg) concentrations based on dietary sources have shown to predict differences in fish; however, they are usually applied at an individual scale and are rarely directed at a known trophic transfer. We combined gut content analysis and stable isotope analysis (δ15N and δ13C) to provide a quantitative estimate of Hg and selenium (Se) biomagnification in the striped marlin (Kajikia audax) and blue marlin (Makaira nigricans) food web from the southwestern Gulf of California. Hg and Se concentrations (mean ± SD; µg g-1, dw) were different among K. audax (Hg = 3.6 ± 2.1, Se = 5.5 ± 5.4) and M. nigricans (Hg = 19.0 ± 29.6, Se = 8.8 ± 10.5). Such variations of element concentrations could be linked to predation with different Hg and Se contents. Diet data presented as prey weight (%W) indicated a higher proportion of large prey fish for the blue marlin than the striped marlin. δ15N and δ13C indicated pelagic food sources with epipelagic preferences for the blue marlin and mesopelagic for the striped marlin. The relationship between Hg concentrations and δ15N was positive along the food web of both marlin species, indicating biomagnification of Hg. However, Se biomagnification was not clearly evidenced, and Se:Hg ratios decreased with δ15N, attributed to increasing Hg concentrations with increased trophic level.

Evidence for Interrupted Biomagnification of Cadmium in Billfish Food Chain Based on Stable Carbon and Nitrogen Isotopes from Southwestern of Gulf of California.

We report cadmium (Cd) concentrations in muscle, liver, and blood of striped marlin (Kajikia audax) and blue marlin (Makaira nigricans), caught in the southwest of the Gulf of California. The average Cd concentration was higher in the liver followed by blood and muscle in descending order. This pattern of Cd concentration may be related to the differences in the physiological function of these tissues. In K. audax, the Cd concentration in muscle and liver increased proportionally with body size, but only in animals that have reached the body size corresponding to first sexual maturity (animals exceeding 155 cm of postorbital length). Interspecific differences in dietary composition and Cd content evidenced that food preferences have a significant effect on the bioaccumulation of Cd. No evidence of Cd biomagnification (progressive bioaccumulation of an element along the food web) was found, as the correlation between logarithmic Cd concentrations and δ15N values was not significant when both billfish and their prey items were included in the calculations. Furthermore, the calculated biotransference factor (transfer of an element from food to consumer) suggested that Cd transference is interrupted from prey to marlins.

Potential medium-term impacts of climate change on tuna and billfish in the Gulf of Mexico: A qualitative framework for management and conservation.

A systematic review of scientific papers on the potential impacts of climate-driven environmental changes on tuna and billfish in the Gulf of Mexico (GOM) was conducted to identify the climate-driven pressures and their associated potential impacts on the reproductive success and survival of tuna and billfish, and which of those impacts may have more relevance for their management and conservation in the GOM by 2050. An Impact Screening Analysis (ISA) was developed to evaluate the potential climate impacts discovered in the literature synthesis by assessing each impact against four criteria, and assigning it a ranking based on likelihood of occurrence (High, Medium, or Low). Results show three types of climate-driven pressures within the High ranking: increased water temperature; changes in ocean circulation and eddy kinetic energy; and changes in storm and wind patterns. Our findings provide valuable information to advance our understanding of key climate-driven physico-chemical processes that can impact the biology of tuna and billfish in the GOM, and enhance conservation and management of these species.

Environmental influences and ontogenetic differences in vertical habitat use of black marlin (Istiompax indica) in the southwestern Pacific.

The black marlin (Istiompax indica) is a highly migratory billfish that occupies waters throughout the tropical and subtropical Indo-Pacific. To characterize the vertical habitat use of I. indica, we examined the temperature-depth profiles collected using 102 pop-up satellite archival tags deployed off the east coast of Australia. Modelling of environmental variables revealed location, sea-surface height deviation, mixed layer depth and dissolved oxygen to all be significant predictors of vertical habitat use. Distinct differences in diel movements were observed between the size classes, with larger size classes of marlin (greater than 50 kg) undertaking predictable bounce-diving activity during daylight hours, while diving behaviour of the smallest size class occurred randomly during both day and night. Overall, larger size classes of I. indica were found to use an increased thermal range and spend more time in waters below 150 m than fish of smaller size classes. The differences in the diving behaviour among size classes were suggested to reflect ontogenetic differences in foraging behaviour or physiology. The findings of this study demonstrate, for the first time to our knowledge, ontogenetic differences in vertical habitat in a species of billfish, and further the understanding of pelagic fish ecophysiology in the presence of global environmental change.

The complete validated mitochondrial genome of the black marlin Istiompax indica (Cuvier, 1832).

Two complete mitochondrial genomes of the black marlin Istiompax indica were assembled from approximately 3.5 and 2.5 million reads produced by Ion Torrent next generation sequencing. The complete genomes were 16,531 bp and 16,532 bp in length consisting of 2 rRNA, 13 protein-coding genes, 22tRNA and 2 coding regions. They demonstrated a similar A + T base (52.6%) to other teleosts. Intraspecific sequence variation was 99.5% for three I. indica mitogenomes and 99.7% for X. gladius. A lower value (85%) was found for the I. platypterus mitogenomes from genbank and accredited to inadvertent inclusion of gene regions from a con-familial species in one record, highlighting the need for cautious downstream use of genbank data.

GEOGRAPHIC DISTRIBUTION OF MOLECULAR VARIANCE WITHIN THE BLUE MARLIN (MAKAIRA NIGRICANS): A HIERARCHICAL ANALYSIS OF ALLOZYME, SINGLE-COPY NUCLEAR DNA, AND MITOCHONDRIAL DNA MARKERS.

This study presents a comparative hierarchical analysis of variance applied to three classes of molecular markers within the blue marlin (Makaira nigricans). Results are reported from analyses of four polymorphic allozyme loci, four polymorphic anonymously chosen single-copy nuclear DNA (scnDNA) loci, and previously reported restriction fragment length polymorphisms (RFLPs) of mitochondrial DNA (mtDNA). Samples were collected within and among the Atlantic and Pacific Oceans over a period of several years. Although moderate levels of genetic variation were detected at both polymorphic allozyme (H = 0.30) and scnDNA loci (H = 0.37), mtDNA markers were much more diverse (h = 0.85). Allele frequencies were significantly different between Atlantic and Pacific Ocean samples at three of four allozyme loci and three of four scnDNA loci. Estimates of allozyme genetic differentiation (θO ) ranged from 0.00 to 0.15, with a mean of 0.08. The θO values for scnDNA loci were similar to those of allozymes, ranging from 0.00 to 0.12 with a mean of 0.09. MtDNA RFLP divergence between oceans (θO = 0.39) was significantly greater than divergence detected at nuclear loci (95% nuclear confidence interval = 0.04-0.11). The fourfold smaller effective population size of mtDNA and male-mediated gene flow may account for the difference observed between nuclear and mitochondrial divergence estimates.