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In November 2020, we observed several individuals and collected one juvenile of an unidentified anthiadine fish (Serranidae) between depths of 250 and 307 m near vertical walls of rocky reefs in the northern Red Sea. Further morphological and molecular analyses revealed that the collected specimen matches Sacura boulengeri, a species previously reported only from the Gulf of Oman to India.

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., [Significance] Global vessel traffic is increasing alongside world economic growth. The potential for rising lethal ship strikes on endangered species of marine megafauna, such as the plankton-feeding whale shark, remains poorly understood since areas of highest overlap are seldom determined across an entire species range. Here we show how satellite tracking whale sharks and large vessel movements globally provides a means to localize high-overlap areas and to determine how collision risk changes in time. Our results point to potential high levels of undetected or unreported ship strikes, which may explain why whale shark populations continue to decline despite protection and low fishing-induced mortality. Collision mitigations in high-collision-risk areas appear necessary to help conserve this iconic species., Funding for data analysis was provided by the UK Natural Environment Research Council (NERC) through a University of Southampton INSPIRE DTP PhD Studentship to F.C.W. Additional funding for data analysis was provided by NERC Discovery Science (NE/R00997/X/1) and the European Research Council (ERC-AdG-2019 883583 OCEAN DEOXYFISH) to D.W.S., Fundação para a Ciência e a Tecnologia (FCT) under PTDC/BIA/28855/2017 and COMPETE POCI-01–0145-FEDER-028855, and MARINFO–NORTE-01–0145-FEDER-000031 (funded by Norte Portugal Regional Operational Program [NORTE2020] under the PORTUGAL 2020 Partnership Agreement, through the European Regional Development Fund–ERDF) to N.Q. FCT also supported N.Q. (CEECIND/02857/2018) and M.V. (PTDC/BIA-COM/28855/2017). D.W.S. was supported by a Marine Biological Association Senior Research Fellowship.

REPLYING TO H. Murua et al. Nature https://doi.org/10.1038/s41586-021-03396-4 (2021) Our previously published paper1 provided global fine-scale spatiotemporal estimates (1° × 1°; monthly) of overlap and fishing exposure risk (FEI) between satellite-tracked shark space use and automatic identification system (AIS) longline fishing effort. We did not assess shark mortality directly, but in addition to replying to the Comment by Murua et al.2, we confirm—using regression analysis of spatially matched data—that fishing-induced pelagic shark mortality (catch per unit effort (CPUE)) is greater where FEI is higher. We focused on assessing shark horizontal spatiotemporal overlap and exposure risk with fisheries because spatial overlap is a major driver of fishing capture susceptibility and previous shark ecological risk assessments (ERAs) assumed a homogenous shark density within species-range distributions3,4,5 or used coarse-scale modelled occurrence data, rather than more ecologically realistic risk estimates in heterogeneous habitats that were selected by sharks over time. Furthermore, our shark spatial exposure risk implicitly accounts for other susceptibility factors with equal or similar probabilities to those commonly used in shark ERAs3,5.

REPLYING TO A. V. Harry & J. M. Braccini Nature https://doi.org/10.1038/s41586-021-03463-w (2021) Our global analysis1 estimated the overlap and fishing exposure risk (FEI) using the space use of satellite-tracked sharks and longline fishing effort monitored by the automatic identification system (AIS). In the accompanying Comment, Harry and Braccini2 draw attention to two localized shark–longline vessel overlap hotspots in Australian waters, stating that 47 fishing vessels were misclassified as longline and purse seine vessels in the Global Fishing Watch (GFW)3 2012–2016 AIS fishing effort data product that we used. This, they propose2, results in misidentifications that highlight fishing exposure hotspots that are subject to an unexpected level of sensitivity in the analysis and they suggest that misidentifications could broadly affect the calculations of fishing exposure and the central conclusions of our study1. We acknowledged in our previously published paper1 that gear reclassifications were likely to occur for a small percentage of the more than 70,000 vessels studied, however, here we demonstrate that even using much larger numbers of vessel reclassifications than those proposed by Harry and Braccini2, the central results and conclusions of our paper1 do not change.

An aggregation of sexually immature whale sharks occurs at a coastal submerged reef near the Saudi Arabian Red Sea coast each spring. We tested the hypothesis that these megaplanktivores become attracted to a prey biomass peak coinciding with their aggregation. Acoustic backscatter of the water column at 120 kHz and 333 kHz –a proxy for potential prey biomass – was continuously measured spanning the period prior to, during, and subsequent to the seasonal whale shark aggregations. No peak in acoustic backscatter was observed at the time of the aggregation. However, we observed a decrease in acoustic backscatter in the last days of deployment, which coincided the trailing end of whale shark season. Organisms forming the main scattering layer performed inverse diel vertical migration, with backscatter peaking at mid-depths during the day and in the deeper half of the water column at night. Target strength analyses suggested the backscatter was likely composed of fish larvae. Subsurface foraging behavior of the whale sharks within this aggregation has not been described, yet this study does not support the hypothesis that seasonal peaks in local whale shark abundance correspond to similar peaks in prey availability.

An Amendment to this paper has been published and can be accessed via a link at the top of the paper.

Fishing has had a profound impact on global reef shark populations, and the absence or presence of sharks is strongly correlated with national socio-economic conditions and reef governance. Decades of overexploitation have devastated shark populations, leaving considerable doubt as to their ecological status(1,2). Yet much of what is known about sharks has been inferred from catch records in industrial fisheries, whereas far less information is available about sharks that live in coastal habitats(3). Here we address this knowledge gap using data from more than 15,000 standardized baited remote underwater video stations that were deployed on 371 reefs in 58 nations to estimate the conservation status of reef sharks globally. Our results reveal the profound impact that fishing has had on reef shark populations: we observed no sharks on almost 20% of the surveyed reefs. Reef sharks were almost completely absent from reefs in several nations, and shark depletion was strongly related to socio-economic conditions such as the size and proximity of the nearest market, poor governance and the density of the human population. However, opportunities for the conservation of reef sharks remain: shark sanctuaries, closed areas, catch limits and an absence of gillnets and longlines were associated with a substantially higher relative abundance of reef sharks. These results reveal several policy pathways for the restoration and management of reef shark populations, from direct top-down management of fishing to indirect improvement of governance conditions. Reef shark populations will only have a high chance of recovery by engaging key socio-economic aspects of tropical fisheries.

The presence of whale sharks Rhincodon typus were recorded around Shib Habil, a small, coastal reef off the Red Sea coast of Saudi Arabia, from 2010 to 2015. A total of 267 suitable photographs resulting in the identification of 136 individuals, were documented from 305 encounters. Sharks were divided evenly between the sexes with no evidence of temporal or spatial segregation. All individuals were immature based on size estimates and, for males, juvenile clasper morphology. Scars were reported for 57% of R. typus with 15% showing evidence of propeller trauma. Estimates of population size and patterns of residency were calculated by modelling the lagged identification rate. Multiple models were run simultaneously and compared using the Akaike information criterion. An open population model was found to best represent the data and estimates a daily abundance between 15 and 34 R. typus during the aggregation season, with local residence times ranging from 4 to 44 days. Residence times away from Shib Habil range from 15 to 156 days with a permanent emigration-death rate between 0·07 and 0·58 individuals year(-1) . These results are broadly similar to those from other aggregations of R. typus, although the observed sexual parity and integration found at this site is unique for the species and needs further study.

Whale sharks (Rhincodon typus) are typically dispersed throughout their circumtropical range, but the species is also known to aggregate in specific coastal areas. Accurate site descriptions associated with these aggregations are essential for the conservation of R. typus, an Endangered species. Although aggregations have become valuable hubs for research, most site descriptions rely heavily on sightings data. In the present study, visual census, passive acoustic monitoring, and long range satellite telemetry were combined to track the movements of R. typus from Shib Habil, a reef-associated aggregation site in the Red Sea. An array of 63 receiver stations was used to record the presence of 84 acoustically tagged sharks (35 females, 37 males, 12 undetermined) from April 2010 to May 2016. Over the same period, identification photos were taken for 76 of these tagged individuals and 38 were fitted with satellite transmitters. In total of 37,461 acoustic detections, 210 visual encounters, and 33 satellite tracks were analyzed to describe the sharks’ movement ecology. The results demonstrate that the aggregation is seasonal, mostly concentrated on the exposed side of Shib Habil, and seems to attract sharks of both sexes in roughly equal numbers. The combined methodologies also tracked 15 interannual homing-migrations, demonstrating that many sharks leave the area before returning in later years. When compared to acoustic studies from other aggregations, these results demonstrate that R. typus exhibits diverse, site-specific ecologies across its range. Sightings-independent data from acoustic telemetry and other sources are an effective means of validating more common visual surveys.

Effective ocean management and the conservation of highly migratory species depend on resolving the overlap between animal movements and distributions, and fishing effort. However, this information is lacking at a global scale. Here we show, using a big-data approach that combines satellite-tracked movements of pelagic sharks and global fishing fleets, that 24% of the mean monthly space used by sharks falls under the footprint of pelagic longline fisheries. Space-use hotspots of commercially valuable sharks and of internationally protected species had the highest overlap with longlines (up to 76% and 64%, respectively), and were also associated with significant increases in fishing effort. We conclude that pelagic sharks have limited spatial refuge from current levels of fishing effort in marine areas beyond national jurisdictions (the high seas). Our results demonstrate an urgent need for conservation and management measures at high-seas hotspots of shark space use, and highlight the potential of simultaneous satellite surveillance of megafauna and fishers as a tool for near-real-time, dynamic management., Data analysis was funded in part by the Marine Biological Association (MBA) and the UK Natural Environment Research Council (NERC) (NE/R00997X/1) (to D.W.S.) with additional research support from the Save Our Seas Foundation and the NERC Oceans 2025 Strategic Research Programme, in which D.W.S. was a principal investigator. D.W.S. was supported by an MBA Senior Research Fellowship, N.Q. by European Regional Development Fund (FEDER) via the Programa Operacional Competitividade e Internacionalização (COMPETE), National Funds via Fundação para a Ciência e a Tecnologia (FCT) under PTDC/MAR/100345/2008 and COMPETE FCOMP-01-0124-FEDER-010580 (to N.Q. and D.W.S.), and Norte Portugal Regional Operational Programme (NORTE 2020) under the PORTUGAL 2020 Partnership Agreement through the European Regional Development Fund (ERDF) in project MarInfo (NORTE-01-0145-FEDER-000031). Additional support was provided by an FCT Investigator Fellowship IF/01611/2013 (N.Q.), FCT Doctoral Fellowship PD/BD/52603/2014 (M.V.), PTDC/MAR-BIO/4458/2012, Xunta de Galicia - Isabel Barreto Program 2009-2012 (G.M.), Australian Research Council (ARC) grant DE170100841 and operational funds from the Australian Institute for Marine Science (AIMS) (both to A.M.M.S.).

The Red Sea has long been recognized as a region of high biodiversity and endemism. Despite this diversity and early history of scientific work, our understanding of the ecology of coral reefs in the Red Sea has lagged behind that of other large coral reef systems. We carried out a quantitative assessment of ISI-listed research published from the Red Sea in eight specific topics (apex predators, connectivity, coral bleaching, coral reproductive biology, herbivory, marine protected areas, non-coral invertebrates and reef-associated bacteria) and compared the amount of research conducted in the Red Sea to that from Australia’s Great Barrier Reef (GBR) and the Caribbean. On average, for these eight topics, the Red Sea had 1/6th the amount of research compared to the GBR and about 1/8th the amount of the Caribbean. Further, more than 50 % of the published research from the Red Sea originated from the Gulf of Aqaba, a small area (

A healthy herbivore community is critical for the ability of a reef to resist and recover from severe disturbances and to regain lost coral cover (i.e., resilience). The densities of the two major herbivorous fish groups (the family Acanthuridae and scarine labrids) were comparatively studied for an inshore reef that was severely impacted by a mass coral bleaching event in 2010 and an unaffected reef within the same region. Densities were found to be significantly higher on the affected reef, most likely due to the high algal densities on that reef. However, densities of herbivores on both reefs were found to be on average about 1–2 orders of magnitude lower than previously published reports from some Pacific reefs and from Red Sea reefs in the Gulf of Aqaba and only slightly higher than Caribbean reefs. Thus, it is predicted that recovery for this reef and similarly affected reefs may be very slow. The protection of herbivores from overfishing and the introduction of other management strategies that maximize reef resilience in Saudi Arabian waters are highly recommended.

(2011). First record of the Pigeye Shark, Carcharhinus amboinensis (Muller & Henle, 1839) (Carcharhiniformes: Carcharhinidae), in the Red Sea. Zoology in the Middle East: Vol. 52, No. 1, pp. 118-121.

Background Off Mafia Island, Tanzania, whale sharks are seen in a small bay that also boasts one of the country's largest coastal fisheries. We set out to determine the population structure of whale sharks here, examine their feeding behaviour and residency, and investigate the interaction between the sharks and fishers. Approach We conducted 261 boat-based surveys at Mafia Island between Oct 2012 and Dec 2015, recording all whale shark sightings as well as counting the numbers of tourist and fishing boats in the survey area during each trip. We estimated shark size visually and with laser photogrammetry and recorded sex, scars and behaviour of each shark. To examine residency, we deployed 67 acoustic tags on whale sharks and 19 receiver stations within the bay, and analysed recapture probability (both visual- and acoustic-based) with generalised additive mixing models. To investigate feeding plankton biomass and community structure, we collected 20 plankton samples when whale sharks were feeding and 202 background samples for comparison. We interviewed fishers and they recorded their fishing locations using a GPS. Results We identified 129 different whale sharks at Mafia Island, 54% of which were resighted in two or more years. Tagged sharks were acoustically detected throughout the year even though sightings were largely limited to Oct-Mar. Sharks ranged in total length from 2.5–9.7 m, were mostly male (87%), and were usually seen feeding (75%) at or near the surface. Dense patches (25 mg m–3) of large sergestids Lucifer hanseni prompted feeding, while background plankton tows were dominated by small copepods (2.5 mg m–3). Small and medium-sized planktivorous fishes also fed on the same patches of sergestids, often together with whale sharks. These smaller species were the main target of the local fishers. Fishing boats operated in the survey area on 90% of the time, but also made catches further afield. Fishers reported a conflict between whale sharks and their fishing operations on 36% of the time, although sharks were usually released from the nets without injury. A mean of 2.6 tourist boats per day searched the same survey area for whale sharks. Conclusions This small, resident group of whale sharks is routinely exposed to tourists, boat traffic and high fishing pressure. Whale sharks are commonly encircled in fishing nets and released, but scars indicate that they do occasionally get injured during this process. Mortalities are rare, based on interviews with fishers and long-term observers. Whale sharks at Mafia Island are unusually resident and face clear local threats. No defined management measures are currently in place, but we present some potential strategies for mitigating threats to whale sharks here.

Background The Red Sea has received little study when compared to other areas. This holds true for sharks in general and for whale sharks in particular. The discovery of a seasonal aggregation off the Saudi Arabian coast has made the targeted study of Red Sea whale sharks more feasible. Assessing the size and structure of this aggregation is a first step toward understanding the basin-scale population within the Red Sea and how that population fits into the wider Indian Ocean. Approach Photo-ID records were collected from 2010 through 2015. These were analyzed and fit to several population models. The most parsimonious model was then used to describe the aggregation. Demographic data, including the size and sex of individual sharks was collected and compared to reports from other aggregations. Size was visually estimated to the nearest half-meter and sex was determined by the presence or absence of claspers between the pelvic fins. Results From 2010 to 2015, 305 encounters with whale sharks were recorded within the aggregation. From these, 267 suitable photos were used to identify 136 individuals. Sharks were divided evenly between the sexes and the distribution of sightings showed no evidence of sexual segregation. All individuals were immature based on size estimates and, in males, juvenile clasper morphology. An open population model was found to best represent the data and estimated a daily abundance between 15 and 34 whale sharks during the aggregation season, with local residence times ranging from 4 to 44 days. Residence times away from Shib Habil ranged from 15 to 156 days with a permanent emigration/mortality rate between 0.07 and 0.58 individuals per year. Conclusions These results are broadly similar to those from other aggregations of R. typus, although the observed sexual parity and integration found at this site are unique for the species and need further study.

Background A recent global study of whale shark population genetics has allowed for better understanding of genetic connections between aggregations in both the Indo-Pacific and Atlantic. This overview included an aggregation found within the Red Sea near Al Lith, Saudi Arabia, however the Mafia Island, Tanzania, aggregation was not part of the study. The ecological behavior of these aggregations differs with the Saudi Arabian individuals showing strong seasonality, while acoustic telemetry data revealed cryptic residency at Mafia Island. Approach Genetic analysis using 11 microsatellite markers was performed on whale sharks from both locations. A combination of primers sourced from previous studies and newly designed primers were used to compare both aggregations and the individuals within. The Red Sea population was compared between 5 seasons spanning 6 years from 2010–2015. The Tanzanian population was compared for 2 field seasons from 2012–2014. Temporal genetic diversity was examined using allelic richness on only the Saudi Arabian individuals due to a short sampling period in Tanzania. Kinship for both aggregations was tested using COLONY and KINALYZER. Results Over a 6 year period, genetic diversity in the Red Sea showed no significant change. Contrasting to other whale shark aggregations, allelic richness in the Red Sea shows no sign of reduction. Kinship analysis using COLONY found two potential sibling pairs in Tanzania. One pair had a high probability (.993) of being a full sibling dyad while the other had a lower probability (.357). There were no sibling pairs identified in the Red Sea. Conclusions The lack of significant change of genetic diversity in Al Lith, Saudi Arabia, differs from a trend at Ningaloo Reef, Australia that showed a decrease in genetic diversity. Although these differences could be driven by location, this should encourage further long term genetic sampling at aggregations to better understand whale shark population trends. The potential of sibling pairs being found within one aggregation warrants further investigation into kinship within and between aggregations throughout the Indo-Pacific.

Although whale sharks ( Rhincodon typus ) have been documented to move thousands of kilometres, they are most frequently observed at a few predictable seasonal aggregation sites. The absence of sharks at the surface during visual surveys has led to the assumption that sharks disperse to places unknown during the long ‘off-seasons’ at most of these locations. Here we compare 2 years of R. typus visual sighting records from Mafia Island in Tanzania to concurrent acoustic telemetry of tagged individuals. Sightings revealed a clear seasonal pattern with a peak between October and February and no sharks observed at other times. By contrast, acoustic telemetry demonstrated year-round residency of R. typus . The sharks use a different habitat in the off-season, swimming deeper and further away from shore, presumably in response to prey distributions. This behavioural change reduces the sharks' visibility, giving the false impression that they have left the area. We demonstrate, for the first time to our knowledge, year-round residency of unprovisioned, individual R. typus at an aggregation site, and highlight the importance of using multiple techniques to study the movement ecology of marine megafauna.

Large planktivores require high-density prey patches to make feeding energetically viable. This is a major challenge for species living in tropical and subtropical seas, such as whale sharks Rhincodon typus. Here, we characterize zooplankton biomass, size structure and taxonomic composition from whale shark feeding events and background samples at Mafia Island, Tanzania. The majority of whale sharks were feeding (73%, 380 of 524 observations), with the most common behaviour being active surface feeding (87%). We used 20 samples collected from immediately adjacent to feeding sharks and an additional 202 background samples for comparison to show that plankton biomass was ∼10 times higher in patches where whale sharks were feeding (25 vs. 2.6 mg m−3). Taxonomic analyses of samples showed that the large sergestid Lucifer hanseni (∼10 mm) dominated while sharks were feeding, accounting for ∼50% of identified items, while copepods (

Background. The Western Indian Ocean is a globally important region for the whale shark Rhincodon typus, with well-studied coastal aggregation sites in southern Mozambique, Seychelles and Djibouti. Here we present an overview of a new study at Mafia Island, Tanzania. Methods. We monitored whale shark abundances on 103 boat trips from October 2012–March 2013. We also used passive acoustic telemetry (VEMCO® V16 tags) and photographic identification to monitor the residency times and local movements of 29 tagged individuals. Shark sizes were estimated using laser photogrammetry. Results. In total we observed 87 individual sharks with a mean of 5.1 ± 5.2 (± SD) per trip and a peak in December of 8.2 ± 6.3. Total length ranged from 4.1 to 9.7 m and almost all sharks were immature. After boat-based visual observations dropped to zero in March 2013 (with the same ongoing sampling effort), the acoustic array still detected 75% of tagged sharks. Tagged individuals were detected by the acoustic array for 73 ± 40 days on average. They showed a strong site fidelity to a 15 km2 area in the inner part of the bay and then progressively moved offshore at the end of the season, matching a decrease in plankton abundance. Sharks were mostly observed feeding on dense patches of the pelagic shrimp Lucifer hanseni, often in association with planktivorous fishes. Photo IDs from 2007-09 and 2012-13 indicate that a large proportion of the juvenile individuals return to Mafia Island each spring-summer. Conclusion. The size range and gender distribution of whale sharks at Mafia Island is similar to other coastal aggregations in the Indian Ocean, but the relatively high site fidelity and residency time stands in contrast.

Conservation efforts aimed at the whale shark, Rhincodon typus, remain limited by a lack of basic information on most aspects of its ecology, including global population structure, population sizes and movement patterns. Here we report on the movements of 47 Red Sea whale sharks fitted with three types of satellite transmitting tags from 2009-2011. Most of these sharks were tagged at a single aggregation site near Al-Lith, on the central coast of the Saudi Arabian Red Sea. Individuals encountered at this site were all juveniles based on size estimates ranging from 2.5-7 m total length with a sex ratio of approximately 1∶1. All other known aggregation sites for juvenile whale sharks are dominated by males. Results from tagging efforts showed that most individuals remained in the southern Red Sea and that some sharks returned to the same location in subsequent years. Diving data were recorded by 37 tags, revealing frequent deep dives to at least 500 m and as deep as 1360 m. The unique temperature-depth profiles of the Red Sea confirmed that several whale sharks moved out of the Red Sea while tagged. The wide-ranging horizontal movements of these individuals highlight the need for multinational, cooperative efforts to conserve R. typus populations in the Red Sea and Indian Ocean.