Detection and localization of Goliath grouper using their low-frequency pulse sounds.

The goal of this paper is to implement and deploy an automated detector and localization model to locate underwater marine organisms using their low-frequency pulse sounds. This model is based on time difference of arrival (TDOA) and uses a two-stage approach, first, to identify the sound and, second, to localize it. In the first stage, an adaptive matched filter (MF) is designed and implemented to detect and determine the timing of the sound pulses recorded by the hydrophones. The adaptive MF measures the signal and noise levels to determine an adaptive threshold for the pulse detection. In the second stage, the detected sound pulses are fed to a TDOA localization algorithm to compute the locations of the sound source. Despite the uncertainties stemming from various factors that might cause errors in position estimates, it is shown that the errors in source locations are within the dimensions of the array. Further, our method was applied to the localization of Goliath grouper pulse-like calls from a six-hydrophone array. It was revealed that the intrinsic error of the model was about 2 m for an array spanned over 50 m. This method can be used to automatically process large amount of acoustic data and provide a precise description of small scale movements of marine organisms that produce low-frequency sound pulses.

Influence of offshore oil and gas structures on seascape ecological connectivity.

Offshore platforms, subsea pipelines, wells and related fixed structures supporting the oil and gas (O&G) industry are prevalent in oceans across the globe, with many approaching the end of their operational life and requiring decommissioning. Although structures can possess high ecological diversity and productivity, information on how they interact with broader ecological processes remains unclear. Here, we review the current state of knowledge on the role of O&G infrastructure in maintaining, altering or enhancing ecological connectivity with natural marine habitats. There is a paucity of studies on the subject with only 33 papers specifically targeting connectivity and O&G structures, although other studies provide important related information. Evidence for O&G structures facilitating vertical and horizontal seascape connectivity exists for larvae and mobile adult invertebrates, fish and megafauna; including threatened and commercially important species. The degree to which these structures represent a beneficial or detrimental net impact remains unclear, is complex and ultimately needs more research to determine the extent to which natural connectivity networks are conserved, enhanced or disrupted. We discuss the potential impacts of different decommissioning approaches on seascape connectivity and identify, through expert elicitation, critical knowledge gaps that, if addressed, may further inform decision making for the life cycle of O&G infrastructure, with relevance for other industries (e.g. renewables). The most highly ranked critical knowledge gap was a need to understand how O&G structures modify and influence the movement patterns of mobile species and dispersal stages of sessile marine species. Understanding how different decommissioning options affect species survival and movement was also highly ranked, as was understanding the extent to which O&G structures contribute to extending species distributions by providing rest stops, foraging habitat, and stepping stones. These questions could be addressed with further dedicated studies of animal movement in relation to structures using telemetry, molecular techniques and movement models. Our review and these priority questions provide a roadmap for advancing research needed to support evidence-based decision making for decommissioning O&G infrastructure.

Classifying Goliath Grouper (Epinephelus itajara) Behaviors from a Novel, Multi-Sensor Tag.

Inertial measurement unit sensors (IMU; i.e., accelerometer, gyroscope and magnetometer combinations) are frequently fitted to animals to better understand their activity patterns and energy expenditure. Capable of recording hundreds of data points a second, these sensors can quickly produce large datasets that require methods to automate behavioral classification. Here, we describe behaviors derived from a custom-built multi-sensor bio-logging tag attached to Atlantic Goliath grouper (Epinephelus itajara) within a simulated ecosystem. We then compared the performance of two commonly applied machine learning approaches (random forest and support vector machine) to a deep learning approach (convolutional neural network, or CNN) for classifying IMU data from this tag. CNNs are frequently used to recognize activities from IMU data obtained from humans but are less commonly considered for other animals. Thirteen behavioral classes were identified during ethogram development, nine of which were classified. For the conventional machine learning approaches, 187 summary statistics were extracted from the data, including time and frequency domain features. The CNN was fed absolute values obtained from fast Fourier transformations of the raw tri-axial accelerometer, gyroscope and magnetometer channels, with a frequency resolution of 512 data points. Five metrics were used to assess classifier performance; the deep learning approach performed better across all metrics (Sensitivity = 0.962; Specificity = 0.996; F1-score = 0.962; Matthew's Correlation Coefficient = 0.959; Cohen's Kappa = 0.833) than both conventional machine learning approaches. Generally, the random forest performed better than the support vector machine. In some instances, a conventional learning approach yielded a higher performance metric for particular classes (e.g., the random forest had a F1-score of 0.971 for backward swimming compared to 0.955 for the CNN). Deep learning approaches could potentially improve behavioral classification from IMU data, beyond that obtained from conventional machine learning methods.

Genetic evidence supports a range extension for the Brazilian cownose ray Rhinoptera brasiliensis in the western North Atlantic.

We report 24 new records of the Brazilian cownose ray Rhinoptera brasiliensis outside its accepted geographic range. Sequencing of a 442-base pair portion of the mitochondrial NADH dehydrogenase subunit 2 gene for 282 Rhinoptera samples revealed eight records off the east coast of the USA and 16 from the eastern Gulf of Mexico. Both sexes of all life stages were documented in all seasons over multiple years in the Indian River and Lake Worth lagoons, Florida, indicating that their range extends further in the western North Atlantic than previously described.

First insights into the vertical habitat use of the whitespotted eagle ray Aetobatus narinari revealed by pop-up satellite archival tags.

The whitespotted eagle ray Aetobatus narinari is a tropical to warm-temperate benthopelagic batoid that ranges widely throughout the western Atlantic Ocean. Despite conservation concerns for the species, its vertical habitat use and diving behaviour remain unknown. Patterns and drivers in the depth distribution of A. narinari were investigated at two separate locations, the western North Atlantic (Islands of Bermuda) and the eastern Gulf of Mexico (Sarasota, Florida, U.S.A.). Between 2010 and 2014, seven pop-up satellite archival tags were attached to A. narinari using three methods: a through-tail suture, an external tail-band and through-wing attachment. Retention time ranged from 0 to 180 days, with tags attached via the through-tail method retained longest. Tagged rays spent the majority of time (82.85 ± 12.17% S.D.) within the upper 10 m of the water column and, with one exception, no rays travelled deeper than ~26 m. One Bermuda ray recorded a maximum depth of 50.5 m, suggesting that these animals make excursions off the fore-reef slope of the Bermuda Platform. Individuals occupied deeper depths (7.42 ± 3.99 m S.D.) during the day versus night (4.90 ± 2.89 m S.D.), which may be explained by foraging and/or predator avoidance. Each individual experienced a significant difference in depth and temperature distributions over the diel cycle. There was evidence that mean hourly depth was best described by location and individual variation using a generalized additive mixed model approach. This is the first study to compare depth distributions of A. narinari from different locations and describe the thermal habitat for this species. Our study highlights the importance of region in describing A. narinari depth use, which may be relevant when developing management plans, whilst demonstrating that diel patterns appear to hold across individuals.

Discerning the dietary habits of the smooth butterfly ray Gymnura lessae using two distinct methods, otolith identification and metagenetics.

Two different methods, metagenetics and free-otolith identification, were used to identify prey in the stomach contents of 531 Gymnura lessae captured by trawling in Mobile Bay, Alabama 2016-2018. Both methods were found to produce analogous results and were therefore combined into a single complete dataset. All prey were teleosts; the families Sciaenidae and Engraulidae were the most important prey (prey specie index of relative importance 89.3% IPSRI ). Multivariate analyses indicated that the diet of G. lessae varied with sex and seasonality. Specifically, variability was probably due to morphologically larger females consuming larger teleost prey species compared with males, whereas seasonal variability was probably due to changes in the available prey community composition. The findings indicate that both metagenetics and free otolith identification, used independently or complementarily, offer robust means of characterising dietary habits for teleost-specialised species such as G. lessae, which may play an important role in the structure and maintenance of coastal food webs such as those in Mobile Bay.

Reproductive physiology of bonefishes (Albula spp.) across the Northwest Bahamas.

Bonefishes (Albula spp.) are classified within the superorder Elopomorpha, which is comprised of over 1000 species that share a unique leptocephalus larval stage. Bonefishes have a circum-tropical distribution, inhabiting inshore shallow water flats and gathering in presumptive nearshore pre-spawn aggregations (PSA) during spawning months. These fishes support economically important recreational fisheries and subsistence fisheries throughout their ranges, yet little is known regarding their reproductive biology. Analysis of oocyte development and nutrient composition, and sex and gonadotrophic hormone levels, was conducted on females sampled in Grand Bahama, Central Andros, and South Andros, The Bahamas, to assess their reproductive state. Fish collected from the flats habitats along all three islands exhibited four major reproductive phases (immature, developing, spawning capable, and regressing). In contrast, all females captured at presumptive PSA sites had eggs in the final stage of oocyte maturation, significantly higher levels of all reproductive hormones (17ß-estradiol, testosterone, and LH), larger vitellogenic oocytes, and oocytes exhibiting germinal vesicle migration and germinal vesicle breakdown. In addition, monthly variability in hormone levels and spawning readiness between Grand Bahama and Andros PSAs suggest that peak spawning times may differ among regions. Fatty acid and free amino acid composition and profiles, with high proportions of docosahexaenoic acid, histidine, and taurine, suggest that these nutrients are not only relevant as energy reserves, but also help achieve buoyancy and osmoregulation of oocytes. This study expands upon our understanding of fish reproductive and developmental physiology, and indicates potential factors influencing the survival and recruitment of bonefishes.

Detection of numerous phycotoxins in young bull sharks (Carcharhinus leucas) collected from an estuary of national significance.

Florida's Indian River Lagoon (IRL) has experienced large-scale, frequent blooms of toxic harmful algae in recent decades. Sentinel, or indicator, species can provide an integrated picture of contaminants in the environment and may be useful to understanding phycotoxin prevalence in the IRL. This study evaluated the presence of phycotoxins in the IRL ecosystem by using the bull shark (Carcharhinus leucas) as a sentinel species. Concentrations of phycotoxins were measured in samples collected from 50 immature bull sharks captured in the IRL between 2018 and 2020. Ultra-performance liquid chromatography/tandem mass spectrometry was used to measure toxins in shark gut contents, plasma, and liver. Analysis of samples (n = 123) demonstrated the presence of multiple phycotoxins (microcystin, nodularin, teleocidin, cylindrospermopsin, domoic acid, okadaic acid, and brevetoxin) in 82 % of sampled bull sharks. However, most detected toxins were in low prevalence (≤25 % of samples, per sample type). This study provides valuable baseline information on presence of multiple phycotoxins in a species occupying a high trophic position in this estuary of national significance.

Visitation patterns of two ray mesopredators at shellfish aquaculture leases in the Indian River Lagoon, Florida.

The Indian River Lagoon is a primary location of field-based "grow-out" for bivalve shellfish aquaculture along Florida's Atlantic coast. Grow-out locations have substantially higher clam densities than surrounding ambient sediment, potentially attracting mollusk predators to the area. Inspired by clammer reports of damaged grow-out gear, we used passive acoustic telemetry to examine the potential interactions between two highly mobile invertivores-whitespotted eagle rays (Aetobatus narinari) and cownose rays (Rhinoptera spp.)-and two clam lease sites in Sebastian, FL and compared these to nearby reference sites (Saint Sebastian River mouth, Sebastian Inlet) from 01 June 2017 to 31 May 2019. Clam lease detections accounted for 11.3% and 5.6% of total detections within the study period, for cownose and whitespotted eagle rays, respectively. Overall, the inlet sites logged the highest proportion of detections for whitespotted eagle rays (85.6%), while cownose rays (11.1%) did not use the inlet region extensively. However, both species had significantly more detections at the inlet receivers during the day, and on the lagoon receivers during the night. Both species exhibited long duration visits (> 17.1 min) to clam lease sites, with the longest visit being 387.5 min. These visit durations did not vary substantially between species, although there was individual variability. Based on generalized additive mixed models, longer visits were observed around 1000 and 1800 h for cownose and whitespotted eagle rays, respectively. Since 84% of all visits were from whitespotted eagle rays and these longer visits were significantly longer at night, this information suggests that observed interactions with the clam leases are potentially underestimated, given most clamming operations occur during daytime (i.e., morning). These results justify the need for continued monitoring of mobile invertivores in the region, including additional experimentation to assess behaviors (e.g., foraging) exhibited at the clam lease sites.

Biophysical larval dispersal models of observed bonefish (Albula vulpes) spawning events in Abaco, The Bahamas: An assessment of population connectivity and ocean dynamics.

Biophysical models are a powerful tool for assessing population connectivity of marine organisms that broadcast spawn. Albula vulpes is a species of bonefish that is an economically and culturally important sportfish found throughout the Caribbean and that exhibits genetic connectivity among geographically distant populations. We created ontogenetically relevant biophysical models for bonefish larval dispersal based upon multiple observed spawning events in Abaco, The Bahamas in 2013, 2018, and 2019. Biological parameterizations were informed through active acoustic telemetry, CTD casts, captive larval rearing, and field collections of related albulids and anguillids. Ocean conditions were derived from the Regional Navy Coastal Ocean Model American Seas dataset. Each spawning event was simulated 100 times using the program Ichthyop. Ten-thousand particles were released at observed and putative spawning locations and were allowed to disperse for the full 71-day pelagic larval duration for A. vulpes. Settlement densities in defined settlement zones were assessed along with interactions with oceanographic features. The prevailing Northern dispersal paradigm exhibited strong connectivity with Grand Bahama, the Berry Islands, Andros, and self-recruitment to lower and upper Abaco. Ephemeral gyres and flow direction within Northwest and Northeast Providence Channels were shown to have important roles in larval retention to the Bahamian Archipelago. Larval development environments for larvae settling upon different islands showed few differences and dispersal was closely associated with the thermocline. Settlement patterns informed the suggestion for expansion of conservation parks in Grand Bahama, Abaco, and Andros, and the creation of a parks in Eleuthera and the Berry Islands to protect fisheries. Further observation of spawning events and the creation of biophysical models will help to maximize protection for bonefish spawning locations and nursery habitat, and may help to predict year-class strength for bonefish stocks throughout the Greater Caribbean.

Diving into the vertical dimension of elasmobranch movement ecology.

Knowledge of the three-dimensional movement patterns of elasmobranchs is vital to understand their ecological roles and exposure to anthropogenic pressures. To date, comparative studies among species at global scales have mostly focused on horizontal movements. Our study addresses the knowledge gap of vertical movements by compiling the first global synthesis of vertical habitat use by elasmobranchs from data obtained by deployment of 989 biotelemetry tags on 38 elasmobranch species. Elasmobranchs displayed high intra- and interspecific variability in vertical movement patterns. Substantial vertical overlap was observed for many epipelagic elasmobranchs, indicating an increased likelihood to display spatial overlap, biologically interact, and share similar risk to anthropogenic threats that vary on a vertical gradient. We highlight the critical next steps toward incorporating vertical movement into global management and monitoring strategies for elasmobranchs, emphasizing the need to address geographic and taxonomic biases in deployments and to concurrently consider both horizontal and vertical movements.

Harmful algal and cyanobacterial toxins in foraging green turtles (Chelonia mydas) in Florida's Big Bend.

Numerous toxin-producing harmful algal (HAB) species occur in Florida's coastal waters. Exposure to these toxins has been shown to have sublethal effects in sea turtles. The objective of this study was to establish concentrations of 10 HAB toxins in plasma samples from green turtles (Chelonia mydas) foraging in Florida's Big Bend. Domoic acid, lyngbyatoxin-A, microcystins, nodularin, and okadaic acid were detected, demonstrating exposure to these HAB toxins, which are also a public health concern.

Movement patterns and habitat use of tiger sharks (Galeocerdo cuvier) across ontogeny in the Gulf of Mexico.

The tiger shark (Galeocerdo cuvier) is globally distributed with established coastal and open-ocean movement patterns in many portions of its range. While all life stages of tiger sharks are known to occur in the Gulf of Mexico (GoM), variability in habitat use and movement patterns over ontogeny have never been quantified in this large marine ecosystem. To address this data gap we fitted 56 tiger sharks with Smart Position and Temperature transmitting tags between 2010 and 2018 and examined seasonal and spatial distribution patterns across the GoM. Additionally, we analyzed overlap of core habitats (i.e., 50% kernel density estimates) among individuals relative to large benthic features (oil and gas platforms, natural banks, bathymetric breaks). Our analyses revealed significant ontogenetic and seasonal differences in distribution patterns as well as across-shelf (i.e., regional) and sex-linked variability in movement rates. Presumably sub-adult and adult sharks achieved significantly higher movement rates and used off-shelf deeper habitats at greater proportions than juvenile sharks, particularly during the fall and winter seasons. Further, female maximum rate of movement was higher than males when accounting for size. Additionally, we found evidence of core regions encompassing the National Oceanographic and Atmospheric Administration designated Habitat Areas of Particular Concern (i.e., shelf-edge banks) during cooler months, particularly by females, as well as 2,504 oil and gas platforms. These data provide a baseline for future assessments of environmental impacts, such as climate variability or oil spills, on tiger shark movements and distribution in the region. Future research may benefit from combining alternative tracking tools, such as acoustic telemetry and genetic approaches, which can facilitate long-term assessment of the species' movement dynamics and better elucidate the ecological significance of the core habitats identified here.

Population connectivity of pelagic megafauna in the Cuba-Mexico-United States triangle.

The timing and extent of international crossings by billfishes, tunas, and sharks in the Cuba-Mexico-United States (U.S.) triangle was investigated using electronic tagging data from eight species that resulted in >22,000 tracking days. Transnational movements of these highly mobile marine predators were pronounced with varying levels of bi- or tri-national population connectivity displayed by each species. Billfishes and tunas moved throughout the Gulf of Mexico and all species investigated (blue marlin, white marlin, Atlantic bluefin tuna, yellowfin tuna) frequently crossed international boundaries and entered the territorial waters of Cuba and/or Mexico. Certain sharks (tiger shark, scalloped hammerhead) displayed prolonged periods of residency in U.S. waters with more limited displacements, while whale sharks and to a lesser degree shortfin mako moved through multiple jurisdictions. The spatial extent of associated movements was generally associated with their differential use of coastal and open ocean pelagic ecosystems. Species with the majority of daily positions in oceanic waters off the continental shelf showed the greatest tendency for transnational movements and typically traveled farther from initial tagging locations. Several species converged on a common seasonal movement pattern between territorial waters of the U.S. (summer) and Mexico (winter).

First insights into the movements of young-of-the-year white sharks (Carcharodon carcharias) in the western North Atlantic Ocean.

In recent years, white sharks (Carcharodon carcharias) have become more accessible to researchers off the northeastern U.S. as feeding aggregation sites have emerged and the population has increased. However, there has been limited research on young-of-the-year (YOY) sharks relative to older age classes in this region. Previous research indicated that YOY white sharks were most frequently observed in the New York Bight, suggesting the region serves a nursery role. To further examine the species' use of this area, we deployed satellite and acoustic tags on ten YOY white sharks (138-166 cm total length) off Long Island, New York. The sharks remained resident in New York Bight waters through summer (August through October), further supporting the notion that the region is a nursery area. Southward movements were observed during fall, with overwintering habitat identified off North and South Carolina shelf waters. Return migrations toward the New York Bight were observed in some individuals the following spring. YOY white sharks in this heavily-populated region are exposed to anthropogenic impacts such as fisheries bycatch and coastal habitat degradation. As juvenile survival rates are important for long-term population sustainability, further research is necessary to assess the potential impacts of these activities on the western North Atlantic white shark population.

An Analysis of Artificial Reef Fish Community Structure along the Northwestern Gulf of Mexico Shelf: Potential Impacts of "Rigs-to-Reefs" Programs.

Artificial structures are the dominant complex marine habitat type along the northwestern Gulf of Mexico (GOM) shelf. These habitats can consist of a variety of materials, but in this region are primarily comprised of active and reefed oil and gas platforms. Despite being established for several decades, the fish communities inhabiting these structures remain poorly investigated. Between 2012 and 2013 we assessed fish communities at 15 sites using remotely operated vehicles (ROVs). Fish assemblages were quantified from standing platforms and an array of artificial reef types (Liberty Ships and partially removed or toppled platforms) distributed over the Texas continental shelf. The depth gradient covered by the surveys (30-84 m) and variability in structure density and relief also permitted analyses of the effects of these characteristics on fish richness, diversity, and assemblage composition. ROVs captured a variety of species inhabiting these reefs from large transient piscivores to small herbivorous reef fishes. While structure type and relief were shown to influence species richness and community structure, major trends in species composition were largely explained by the bottom depth where these structures occurred. We observed a shift in fish communities and relatively high diversity at approximately 60 m bottom depth, confirming trends observed in previous studies of standing platforms. This depth was also correlated with some of the largest Red Snapper captured on supplementary vertical longline surveys. Our work indicates that managers of artificial reefing programs (e.g., Rigs-to-Reefs) in the GOM should carefully consider the ambient environmental conditions when designing reef sites. For the Texas continental shelf, reefing materials at a 50-60 m bottom depth can serve a dual purpose of enhancing diving experiences and providing the best potential habitat for relatively large Red Snapper.

Distribution and dynamic habitat use of young bull sharks Carcharhinus leucas in a highly stratified northern Gulf of Mexico estuary.

Understanding how animals alter habitat use in response to changing abiotic conditions is important for effective conservation management. For bull sharks (Carcharhinus leucas), habitat use has been widely examined in the eastern and western Gulf of Mexico; however, knowledge of their movements and the factors influencing them is lacking for populations in the more temperate north-central Gulf of Mexico. To examine how changes in hydrographic conditions affected the presence of young bull sharks in Mobile Bay, Alabama, thirty-five sharks were fitted with internal acoustic transmitters and monitored with an acoustic monitoring array consisting of thirty-three receivers between June 2009 and December 2010. Tagged sharks ranged in size from 60 to 114 cm fork length and were detected between the upper and lower portions of Mobile Bay. Despite a variety of freshwater sources associated with this highly productive estuary, sharks were most consistently detected at the largest input to the system--the Mobile and Tensaw Rivers. Our findings suggest a combination of hydrographic factors interact to influence the distribution of juvenile bull sharks in Mobile Bay. The factors affecting the probability of detecting at least one bull shark varied both temporally (2009 vs 2010) and spatially (upper vs lower bay). Electivity analysis demonstrated that bull sharks showed highest affinity for warm water (29-32 °C), moderate salinities (10-11 psu) and normoxic waters (5-7 mg/l), although these patterns were not consistent between regions or across years. We suggest future studies coupling telemetry and hydrographic variables should, when possible, consider the interactions of multiple environmental parameters when defining the dynamic factors explaining the spatial distribution of coastal sharks.

Estimating the potential impacts of large mesopredators on benthic resources: integrative assessment of spotted eagle ray foraging ecology in Bermuda.

Declines of large sharks and subsequent release of elasmobranch mesopredators (smaller sharks and rays) may pose problems for marine fisheries management as some mesopredators consume exploitable shellfish species. The spotted eagle ray (Aetobatus narinari) is the most abundant inshore elasmobranch in subtropical Bermuda, but its predatory role remains unexamined despite suspected abundance increases and its hypothesized specialization for mollusks. We utilized a combination of acoustic telemetry, benthic invertebrate sampling, gut content analysis and manipulative experiments to assess the impact of spotted eagle rays on Bermudian shellfish resources. Residency and distribution of adult spotted eagle rays was monitored over two consecutive summers in Harrington Sound (HS), an enclosed inshore lagoon that has historically supported multiple recreational and commercial shellfish species. Telemetered rays exhibited variable fidelity (depending on sex) to HS, though generally selected regions that supported relatively high densities of potential mollusk prey. Gut content analysis from rays collected in HS revealed a diet of mainly bivalves and a few gastropods, with calico clam (Macrocallista maculata) representing the most important prey item. Manipulative field and mesocosm experiments with calico clams suggested that rays selected prey patches based on density, though there was no evidence of rays depleting clam patches to extirpation. Overall, spotted eagle rays had modest impacts on local shellfish populations at current population levels, suggesting a reduced role in transmitting cascading effects from apex predator loss. However, due to the strong degree of coupling between rays and multiple protected mollusks in HS, ecosystem-based management that accounts for ray predation should be adopted.