Linking vertical movements of large pelagic predators with distribution patterns of biomass in the open ocean.

Many predator species make regular excursions from near-surface waters to the twilight (200 to 1,000 m) and midnight (1,000 to 3,000 m) zones of the deep pelagic ocean. While the occurrence of significant vertical movements into the deep ocean has evolved independently across taxonomic groups, the functional role(s) and ecological significance of these movements remain poorly understood. Here, we integrate results from satellite tagging efforts with model predictions of deep prey layers in the North Atlantic Ocean to determine whether prey distributions are correlated with vertical habitat use across 12 species of predators. Using 3D movement data for 344 individuals who traversed nearly 1.5 million km of pelagic ocean in [Formula: see text]42,000 d, we found that nearly every tagged predator frequented the twilight zone and many made regular trips to the midnight zone. Using a predictive model, we found clear alignment of predator depth use with the expected location of deep pelagic prey for at least half of the predator species. We compared high-resolution predator data with shipboard acoustics and selected representative matches that highlight the opportunities and challenges in the analysis and synthesis of these data. While not all observed behavior was consistent with estimated prey availability at depth, our results suggest that deep pelagic biomass likely has high ecological value for a suite of commercially important predators in the open ocean. Careful consideration of the disruption to ecosystem services provided by pelagic food webs is needed before the potential costs and benefits of proceeding with extractive activities in the deep ocean can be evaluated.

Global collision-risk hotspots of marine traffic and the world's largest fish, the whale shark.

Marine traffic is increasing globally yet collisions with endangered megafauna such as whales, sea turtles, and planktivorous sharks go largely undetected or unreported. Collisions leading to mortality can have population-level consequences for endangered species. Hence, identifying simultaneous space use of megafauna and shipping throughout ranges may reveal as-yet-unknown spatial targets requiring conservation. However, global studies tracking megafauna and shipping occurrences are lacking. Here we combine satellite-tracked movements of the whale shark, Rhincodon typus, and vessel activity to show that 92% of sharks' horizontal space use and nearly 50% of vertical space use overlap with persistent large vessel (>300 gross tons) traffic. Collision-risk estimates correlated with reported whale shark mortality from ship strikes, indicating higher mortality in areas with greatest overlap. Hotspots of potential collision risk were evident in all major oceans, predominantly from overlap with cargo and tanker vessels, and were concentrated in gulf regions, where dense traffic co-occurred with seasonal shark movements. Nearly a third of whale shark hotspots overlapped with the highest collision-risk areas, with the last known locations of tracked sharks coinciding with busier shipping routes more often than expected. Depth-recording tags provided evidence for sinking, likely dead, whale sharks, suggesting substantial "cryptic" lethal ship strikes are possible, which could explain why whale shark population declines continue despite international protection and low fishing-induced mortality. Mitigation measures to reduce ship-strike risk should be considered to conserve this species and other ocean giants that are likely experiencing similar impacts from growing global vessel traffic.

Direct measurement of cruising and burst swimming speeds of the shortfin mako shark (Isurus oxyrinchus) with estimates of field metabolic rate.

The shortfin mako shark is a large-bodied pursuit predator thought to be capable of the highest swimming speeds of any elasmobranch and potentially one of the highest energetic demands of any marine fish. Nonetheless, few direct speed measurements have been reported for this species. Here, animal-borne bio-loggers attached to two mako sharks were used to provide direct measurements of swimming speeds, kinematics and thermal physiology. Mean sustained (cruising) speed was 0.90 m s-1 (±0.07 s.d.) with a mean tail-beat frequency (TBF) of 0.51 Hz (±0.16 s.d.). The maximum burst speed recorded was 5.02 m s-1 (TBFmax = 3.65 Hz) from a 2 m long female. Burst swimming was sustained for 14 s (mean speed = 2.38 m s-1 ), leading to a 0.24°C increase in white muscle temperature in the 12.5 min after the burst. Routine field metabolic rate was estimated at 185.2 mg O2 kg-1 h-1 (at 18°C ambient temperature). Gliding behaviour (zero TBF) was more frequently observed after periods of high activity, especially after capture when internal (white muscle) temperature approached 21°C (ambient temperature: 18.3°C), indicating gliding probably functions as an energy recovery mechanism and limits further metabolic heat production. The results show shortfin mako sharks generally cruise at speeds similar to other endothermic fish - but faster than ectothermic sharks - with the maximum recorded burst speed being among the highest so far directly measured among sharks, tunas and billfishes. This newly recorded high-oxygen-demand performance of mako sharks suggests it may be particularly vulnerable to habitat loss due to climate-driven ocean deoxygenation.

Fish biodiversity of Saint Peter and Saint Paul's Archipelago, Mid-Atlantic Ridge, Brazil: new records and a species database.

Saint Peter and Saint Paul's Archipelago (SPSPA), one of the smallest and most isolated island groups in the world, is situated on the Mid-Atlantic Ridge, between Brazil and the African continent. SPSPA has low species richness and high endemism; nonetheless, the diversity of fishes from deep habitats (>30 m depth) had not been previously studied in detail. Several expeditions conducted between 2009 and 2018 explored the shallow and deep reefs of SPSPA using scuba, closed-circuit rebreathers, manned submersibles, baited remote underwater stereo-videos (stereo-BRUV) and fishing between 0 and 1050 m depth. These expeditions yielded 41 new records of fishes for SPSPA: 9 in open waters, 9 in shallow waters (0-30 m), 8 in mesophotic ecosystems (30-150 m) and 15 in deeper reefs (>150 m). Combined with literature records of adult pelagic, shallow and deep-reef species, as well as larvae, the database of the fish biodiversity for SPSPA currently comprises 225 species (169 recorded as adult fishes and 79 as larvae, with 23 species found in both stages). Most of them (112) are pelagic, 86 are reef-associated species and 27 are deep-water specialists. Species accumulation curves show that the number of fish species has not yet reached an asymptote. Whereas the number of species recorded in SPSPA is similar to that in other oceanic islands in the Atlantic Ocean, the proportion of shorefishes is relatively lower, and the endemism level is the third highest in the Atlantic. Twenty-nine species are listed as threatened with extinction. Observations confirm the paucity of top predators on shallow rocky reefs of the island, despite the presence of several pelagic shark species around SPSPA. Because all of the endemic species are reef associated, it is argued that the new marine-protected areas created by the Brazilian government do not ensure the protection and recovery of SPSPA's biodiversity because they allow exploitation of the most vulnerable species around the archipelago itself. This study suggests a ban on reef fish exploitation inside an area delimited by the 1000 m isobath around the islands (where all known endemics are concentrated) as the main conservation strategy to be included in the SPSPA management plan being prepared by the Brazilian government.

Seasonal aggregation and diel activity by the sicklefin devil ray Mobula tarapacana off a small, equatorial outcrop of the Mid-Atlantic Ridge.

The present study aimed to analyse occurrence patterns, relative abundance and habitat use by the sicklefin devil ray Mobula tarapacana in the Saint Peter and Saint Paul Archipelago (SPSPA), Brazil, located in the Mid-Atlantic Ridge. Sampling was conducted between December 2008 and December 2012 by surface surveys and tagging with conventional tags and acoustic transmitters. Mobula tarapacana were sighted in the SPSPA year round but the highest frequency of occurrence was observed from January to June, probably due to greater food availability during this season. However, M. tarapacana might also use the area for reproductive purposes based on evidence of mating behaviour. The residency period of tagged rays measured by three autonomous acoustic receivers deployed around SPSPA spanned from 1 to 74 days with most of the detections occurring during daytime. These results suggest that the SPSPA could be an ecologically-important aggregation area for M. tarapacana, possibly providing the species with a feeding or resting habitat while transiting in the equatorial Atlantic Ocean.

Identifying priority sites for whale shark ship collision management globally.

The expansion of the world's merchant fleet poses a great threat to the ocean's biodiversity. Collisions between ships and marine megafauna can have population-level consequences for vulnerable species. The Endangered whale shark (Rhincodon typus) shares a circumglobal distribution with this expanding fleet and tracking of movement pathways has shown that large vessel collisions pose a major threat to the species. However, it is not yet known whether they are also at risk within aggregation sites, where up to 400 individuals can gather to feed on seasonal bursts of planktonic productivity. These "constellation" sites are of significant ecological, socio-economic and cultural value. Here, through expert elicitation, we gathered information from most known constellation sites for this species across the world (>50 constellations and >13,000 individual whale sharks). We defined the spatial boundaries of these sites and their overlap with shipping traffic. Sites were then ranked based on relative levels of potential collision danger posed to whale sharks in the area. Our results showed that researchers and resource managers may underestimate the threat posed by large ship collisions due to a lack of direct evidence, such as injuries or witness accounts, which are available for other, sub-lethal threat categories. We found that constellations in the Arabian Sea and adjacent waters, the Gulf of Mexico, the Gulf of California, and Southeast and East Asia, had the greatest level of collision threat. We also identified 39 sites where peaks in shipping activity coincided with peak seasonal occurrences of whale sharks, sometimes across several months. Simulated collision mitigation options estimated potentially minimal impact to industry, as most whale shark core habitat areas were small. Given the threat posed by vessel collisions, a coordinated, multi-national approach to mitigation is needed within priority whale shark habitats to ensure collision protection for the species.

Hitchhiking to the abyss.

We investigated, for the first time, the hitchhiker-host fidelity of deep-diving whale sharks and Chilean devil rays. We found that two of the most ubiquitous oceanic hitchhikers, the common remora and the pilot fish, are able to follow their hosts to bathypelagic depths, where they are exposed to extreme gradients of light, dissolved oxygen, temperature, and pressure. We documented a deep dive of a large whale shark hosting remoras and pilot fish. Common remora was observed at the deepest section of the dive, at 1460 m, where the water temperature was 3.6°C. A pilot fish was recorded at 900 m, during the ascent phase, with the water temperature of 7.5°C. Although the adaptations that allow these hitchhikers to mitigate the impacts of such extreme environmental conditions remain unknown, we discuss these findings in the framework of the ecophysiology of deep diving and the hitchhiker-host fidelity.

First insights into the shortfin mako shark (Isurus oxyrinchus) fine-scale swimming behaviour.

As regional endotherms, lamnid sharks can sustain high cruising speeds and perform frequent speed bursts. However, since endothermy comes with high energetic costs, lamnids may adopt different swimming strategies to manage their energy budget. Understanding such strategies is essential to provide behavioural and physiological context to their broader movement ecology. The endangered shortfin mako (Isurus oxyrinchus) possibly has the highest energy requirements among lamnids, but our understanding of its swimming behaviour is still limited. We equipped three shortfin mako sharks with high-resolution multi-sensor tags to measure their swimming kinematics in the wild. While swimming horizontally, individuals favoured tail-beat frequencies around 0.6 Hz at speeds comparable to those of ectothermic sharks (ca 0.5 m s-1). All individuals displayed yo-yo-like diving patterns where, for a given tail-beat frequency, speeds were higher during descents, as expected for a negatively buoyant fish. Contrary to what was expected, gliding was almost absent (less than 1.31%). Speed bursts reaching up to 3.6 m s-1 were observed during the day but ceased shortly after dusk, implying a diel change in swimming behaviour. As large-scale research efforts are hindered by this species' increasing rarity, opportunistic high-resolution datasets, like the present, are fundamental to improve our understanding of shortfin mako's behaviour and ecology.

Depth and temperature preferences of meagre, Argyrosomus regius, as revealed by satellite telemetry.

Argyrosomus regius (commonly referred to as meagre), is one of Europe's largest coastal bony fish species and supports important recreational and commercial fisheries in the Atlantic and Mediterranean coasts. Demand for this species, and more recently for their swim bladders, has led to regional population declines and growing importance as an aquaculture species. Despite intense research in captivity, little is known about the spatial ecology of A. regius's wild population, including basic information such as vertical migrations and depth/temperature preferences. Previous research based on indirect data suggests a seasonal habitat shift from shallow to deeper waters, but this has never been validated through direct high-resolution movement data. In this study, we tagged 13 adult A. regius with pop-up satellite archival tags in the South of Portugal, which successfully returned data from 11 individuals including high-resolution data from six recovered tags (mean, range: 167 days, 28-301 days). We found that adults of this population spend 95.2% of their time between 5 and 75 m depth (mean ± SD, 30.9m ± 18.3m) and do not venture beyond 125 m. Across seasons, A. regius move across water temperatures between 13.3 and 24.8°C with a preferred thermal range between 14 and 18°C where they spent 75.4% of their time. The inferential modelling using this electronic data validated previous hypotheses by showing significant differences between a shallower and warmer summer habitat vs. a deeper and cooler winter habitat. Visual investigation of the diel effects on depth preferences suggests subtle changes in depth use between day and night during the warmer months of the year. We speculate that these patterns are in response to the species' behavioural ecology and physiology, reflecting the seasonal changes in water stratification and presence of prey, as well as on the species reproduction, which results in summer spawning aggregations in shallower areas.

Whale Shark (Rhincodon typus) Seasonal Occurrence, Abundance and Demographic Structure in the Mid-Equatorial Atlantic Ocean.

Whale sharks are generally associated with environmental factors that drive their movements to specific locations where food availability is high. Consequently, foraging is believed to be the main reason for the formation of whale shark aggregations. Feeding aggregations occur mainly in nearshore areas and are composed primarily of immature individuals. Conversely, aggregations of mature adults are rarely observed, and their occurrence is correlated with oceanic environments. Despite an increase in the number of whale shark studies, information on mating and parturition grounds is still lacking. In the present work, we assessed the ecological and behavioural aspects of the whale sharks that visit the archipelago of São Pedro and São Paulo (ASPSP), located ~1,000 km off the coast of Brazil in the equatorial Atlantic Ocean. Forty-nine whale sharks were recorded from February 2005 to May 2014. The estimated mean ± SD size was 8.27 ± 2.52 m (range: 2.5-14.0 m) with no significant differences in size across the year. The maturational stages were classified by size as immature (<8.0 m; 32.56%) and mature (>9.0 m; 46.51%); with almost half of the observed animals being mature specimens. The majority of sightings occurred between February and June. During this period, the ocean current weakens and the waters are enriched by eggs and larvae of fishes and invertebrates that attract marine life to forage. At the same time, evidence of reproductive activity in adult females (i.e. swollen abdomen and bite marks on the pectoral fins), and the potential mating behaviour exhibited by one male, suggest that the ASPSP area might also have a role in whale shark reproduction. Irrespective of its use for feeding or reproduction, this insular habitat serves as a meeting point for both juvenile and adult whale sharks, and may play an important ecological role for the species.