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 vessel 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 potential collision mitigation options estimated a minimal impact to industry, as most whale shark core habitat areas were relatively small. Given the threat posed by vessel collisions, a coordinated, multi-national approach to collision mitigation is needed within priority whale shark habitats to ensure collision protection for the species.

Emergent community architecture despite distinct diversity in the global whale shark (Rhincodon typus) epidermal microbiome.

Microbiomes confer beneficial physiological traits to their host, but microbial diversity is inherently variable, challenging the relationship between microbes and their contribution to host health. Here, we compare the diversity and architectural complexity of the epidermal microbiome from 74 individual whale sharks (Rhincodon typus) across five aggregations globally to determine if network properties may be more indicative of the microbiome-host relationship. On the premise that microbes are expected to exhibit biogeographic patterns globally and that distantly related microbial groups can perform similar functions, we hypothesized that microbiome co-occurrence patterns would occur independently of diversity trends and that keystone microbes would vary across locations. We found that whale shark aggregation was the most important factor in discriminating taxonomic diversity patterns. Further, microbiome network architecture was similar across all aggregations, with degree distributions matching Erdos-Renyi-type networks. The microbiome-derived networks, however, display modularity indicating a definitive microbiome structure on the epidermis of whale sharks. In addition, whale sharks hosted 35 high-quality metagenome assembled genomes (MAGs) of which 25 were present from all sample locations, termed the abundant 'core'. Two main MAG groups formed, defined here as Ecogroup 1 and 2, based on the number of genes present in metabolic pathways, suggesting there are at least two important metabolic niches within the whale shark microbiome. Therefore, while variability in microbiome diversity is high, network structure and core taxa are inherent characteristics of the epidermal microbiome in whale sharks. We suggest the host-microbiome and microbe-microbe interactions that drive the self-assembly of the microbiome help support a functionally redundant abundant core and that network characteristics should be considered when linking microbiomes with host health.

Movement and vertical habitat use of yellowfin tuna Thunnus albacares in a vertically compressed habitat: the Galápagos Marine Reserve.

Tropical pelagic predators are exploited by fisheries and their movements are influenced by factors including prey availability, temperature, and dissolved oxygen levels. As the biophysical parameters vary greatly within the range of circumtropical species, local studies are needed to define those species' habitat preference and model possible behavioral responses under different climate change scenarios. Here, we tagged yellowfin tuna Thunnus albacares in the Galápagos Marine Reserve and tracked the horizontal and vertical movements of eight individuals for 4-97 days. The tuna traveled a mean of 13.6 km day-1 horizontally and dispersed throughout the archipelago and in offshore waters inside the Galápagos Marine Reserve and in the surrounding Ecuadorian exclusive economic zone. Vertically, they traveled a mean of 2 km day-1 , although high-resolution data from a recovered tag suggested that transmitted data underestimated their vertical movement by a factor of 5.5. The tracked yellowfin tuna spent most of their time near the surface, with an overall mean swimming depth of 24.3 ± 46.6 m, and stayed shallower at night (11.1 ± 16.3 m) than during the day ( 37.7 ± 60.9 m), but on occasion dived to cold, oxygen-poor waters below 200 m. Deep dives were commonly made during the day with a mean recovery period of 51 min between exposures to modeled oxygen-limiting conditions <1.5 mL L-1 , presumably to re-oxygenate. The depth and frequency of dives were likely limited by dissolved oxygen levels, as oxygen-depleted conditions reach shallow depths in this region. The main habitat of tracked yellowfin tunas was in the shallow mixed layer, which may leave them vulnerable to fishing. Vertical expansion of low-oxygen waters under future climate change scenarios may further compress their habitat, increasing their vulnerability to surface fishing gear.

Distribution of the reef manta ray Mobula alfredi and the oceanic manta ray Mobula birostris in the Philippines: a collaborative effort for conservation.

Little is known about manta ray population size, structure and connectivity in the Philippines. In collaboration with dive operators, non-governmental organizations and authorities, sightings of manta rays were collated into a single national database. Using in-water photographs and videos gathered through citizen science and dedicated research efforts, this study compiled sightings between 2004 and 2020, showing 22 separate sites throughout the archipelago with manta rays present. A total of 392 individual reef manta rays (Mobula alfredi) and 107 oceanic manta rays (Mobula birostris) were identified from the collected footage. Four specific sites in the provinces of Masbate and Palawan together hosted 89% of all identified individuals and accounted for 95% of sightings, highlighting these areas are key aggregation sites. This study also reports the movements of M. birostris within the Philippines, based on photo-identification of three individuals moving 150 km between Cebu and Masbate. Despite the growing number of recreational divers in Daanbantayan and San Jacinto, an 80% decline in M. birostris sightings was observed at these sites. To ensure effective future conservation, it is recommended that efforts focus on the identification and protection of manta ray hotspots and migratory corridors, the creation of a sustainable tourism framework and, most important, the implementation of mitigation strategies to reduce fisheries interactions.

Fatty acid profiles of more than 470 marine species from the Southern Hemisphere.

Lipid and fatty acid datasets are commonly used to assess the nutritional composition of organisms, trophic ecology, and ecosystem dynamics. Lipids and their fatty acid constituents are essential nutrients to all forms of life because they contribute to biological processes such as energy flow and metabolism. Assessment of total lipids in tissues of organisms provides information on energy allocation and life-history strategies and can be an indicator of nutritional condition. The analysis of an organism's fatty acids is a widely used technique for assessing nutrient and energy transfer, and dietary interactions in food webs. Although there have been many published regional studies that assessed lipid and fatty acid compositions, many only report the mean values of the most abundant fatty acids. There are limited individual records available for wider use in intercomparison or macro-scale studies. This dataset consists of 4856 records of individual and pooled samples of at least 470 different marine consumer species sampled from tropical, temperate, and polar regions around Australia and in the Southern, Indian, and Pacific Oceans from 1989 to 2018. This includes data for a diverse range of taxa (zooplankton, fish, cephalopods, chondrichthyans, and marine mammals), size ranges (0.02 cm to ~13 m), and that cover a broad range of trophic positions (2.0-4.6). When known, we provide a record of species name, date of sampling, sampling location, body size, relative (%) measurements of tissue-specific total lipid content and abundant fatty acids, and absolute content (mg 100 g-1 tissue) of eicosapentaenoic acid (EPA, 20:5n3) and docosahexaenoic acid (DHA, 22:6n3) as important long-chain (≥C20 ) polyunsaturated omega-3 fatty acids. These records form a solid basis for comparative studies that will facilitate a broad understanding of the spatial and temporal distribution of marine lipids globally. The dataset also provides reference data for future dietary assessments of marine predators and model assessments of potential impacts of climate change on the availability of marine lipids and fatty acids. There are 480 data records within our data file for which the providers have requested that permission for reuse be granted, with the likely condition that they are included as a coauthor on the reporting of the dataset. Records with this condition are indicated by a "yes" under "Conditions_of_data_use" in Data S1: Marineconsumer_FAdata.csv (see Table 2 in Metadata S1 for more details). For all other data records marked as "No" under "Conditions_of_data_use," there are no copyright restrictions for research and/or teaching purposes. We request that users acknowledge use of the data in publications, research proposals, websites, and other outlets via formal citation of this work and original data sources as applicable.

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.

Pieces in a global puzzle: Population genetics at two whale shark aggregations in the western Indian Ocean.

The whale shark Rhincodon typus is found throughout the world's tropical and warm-temperate ocean basins. Despite their broad physical distribution, research on the species has been concentrated at a few aggregation sites. Comparing DNA sequences from sharks at different sites can provide a demographically neutral understanding of the whale shark's global ecology. Here, we created genetic profiles for 84 whale sharks from the Saudi Arabian Red Sea and 72 individuals from the coast of Tanzania using a combination of microsatellite and mitochondrial sequences. These two sites, separated by approximately 4500 km (shortest over-water distance), exhibit markedly different population demographics and behavioral ecologies. Eleven microsatellite DNA markers revealed that the two aggregation sites have similar levels of allelic richness and appear to be derived from the same source population. We sequenced the mitochondrial control region to produce multiple global haplotype networks (based on different alignment methodologies) that were broadly similar to each other in terms of population structure but suggested different demographic histories. Data from both microsatellite and mitochondrial markers demonstrated the stability of genetic diversity within the Saudi Arabian aggregation site throughout the sampling period. These results contrast previously measured declines in diversity at Ningaloo Reef, Western Australia. Mapping the geographic distribution of whale shark lineages provides insight into the species' connectivity and can be used to direct management efforts at both local and global scales. Similarly, understanding historical fluctuations in whale shark abundance provides a baseline by which to assess current trends. Continued development of new sequencing methods and the incorporation of genomic data could lead to considerable advances in the scientific understanding of whale shark population ecology and corresponding improvements to conservation policy.

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.

Satellite tracking of rehabilitated sea turtles suggests a high rate of short-term survival following release.

The rehabilitation of wildlife can contribute directly to the conservation of threatened species by helping to maintain wild populations. This study focused on determining the post-rehabilitation survival and spatial ecology of sea turtles and on comparing the movements of individuals with flipper amputations (amputees) to non-amputee animals. Our aims were to assess whether rehabilitated sea turtles survive after release, to compare and contrast the movement characteristics of the different species of sea turtles we tracked, and to examine whether amputees and non-amputees within species behaved similarly post-release. Twenty-six rehabilitated sea turtles from four species, including hawksbill Eretmochelys imbricata (n = 12), loggerhead Caretta caretta (n = 11), green Chelonia mydas (n = 2), and olive ridley Lepidochelys olivacea (n = 1) sea turtles from the United Arab Emirates were fitted with satellite tags before release. Rehabilitation times ranged from 89 to 817 days (mean 353 ± 237 days). Post-release movements and survival were monitored for 8 to 387 days (mean 155 ± 95 days) through satellite tracking. Tag data suggested that three tracked sea turtles died within four days of release, one after 27 days, and one after 192 days from what are thought to be anthropogenic factors unrelated to their pre-rehabilitation ailments. We then compared habitat use and movement characteristics among the different sea turtle species. Although half of all turtles crossed one or more international boundaries, dispersal varied among species. Loggerhead turtles had a high dispersal, with 80% crossing an international boundary, while hawksbill turtles displayed higher post-release residency, with 66% remaining within UAE territorial waters. Amputee turtles moved similarly to non-amputee animals of the same species. Loggerhead turtles travelled faster (mean ± sd = 15.3 ± 8 km/day) than hawksbill turtles (9 ± 7 km/day). Both amputee and non-amputee sea turtles within a species moved similarly. Our tracking results highlight that rehabilitated sea turtles, including amputees, can successfully survive in the wild following release for up to our ~one-year monitoring time therefore supporting the suitability for release of sea turtles that have recovered from major injuries such as amputations. However, more broadly, the high mortality from anthropogenic factors in the Arabian Gulf region is clearly a serious issue and conservation challenge.

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.

DNA metabarcoding assays reveal a diverse prey assemblage for Mobula rays in the Bohol Sea, Philippines.

Diet studies provide base understanding of trophic structure and are a valuable initial step for many fields of marine ecology, including conservation and fisheries biology. Considerable complexity in marine trophic structure can exist due to the presence of highly mobile species with long life spans. Mobula rays are highly mobile, large, planktivorous elasmobranchs that are frequently caught either directly or as bycatch in fisheries, which, combined with their conservative life history strategy, makes their populations susceptible to decline in intensely fished regions. Effective management of these iconic and vulnerable species requires an understanding of the diets that sustain them, which can be difficult to determine using conventional sampling methods. We use three DNA metabarcode assays to identify 44 distinct taxa from the stomachs (n = 101) of four sympatric Mobula ray species (Mobula birostris, Mobula tarapacana, Mobula japanica, and Mobula thurstoni) caught over 3 years (2013-2015) in a direct fishery off Bohol in the Philippines. The diversity and incidence of bony fishes observed in ray diets were unprecedented. Nevertheless, rays showed dietary overlap, with krill (Euphausia) dominating their diet. Our results provide a more detailed assessment of sympatric ray diets than was previously described and reveal the complexity that can exist in food webs at critical foraging habitats.

Satellite tracking of juvenile whale sharks in the Sulu and Bohol Seas, Philippines.

The whale shark Rhincodon typus was uplisted to 'Endangered' in the 2016 IUCN Red List due to >50% population decline, largely caused by continued exploitation in the Indo-Pacific. Though the Philippines protected the whale shark in 1998, concerns remain due to continued take in regional waters. In light of this, understanding the movements of whale sharks in the Philippines, one of the most important hotspots for the species, is vital. We tagged 17 juvenile whale sharks with towed SPOT5 tags from three general areas in the Sulu and Bohol Seas: Panaon Island in Southern Leyte, northern Mindanao, and Tubbataha Reefs Natural Park (TRNP). The sharks all remained in Philippine waters for the duration of tracking (6-126 days, mean 64). Individuals travelled 86-2,580 km (mean 887 km) at a mean horizontal speed of 15.5 ± 13.0 SD km day-1. Whale sharks tagged in Panaon Island and Mindanao remained close to shore but still spent significant time off the shelf (>200 m). Sharks tagged at TRNP spent most of their time offshore in the Sulu Sea. Three of twelve whale sharks tagged in the Bohol Sea moved through to the Sulu Sea, whilst two others moved east through the Surigao Strait to the eastern coast of Leyte. One individual tagged at TRNP moved to northern Palawan, and subsequently to the eastern coast of Mindanao in the Pacific Ocean. Based on inferred relationships with temperature histograms, whale sharks performed most deep dives (>200 m) during the night, in contrast to results from whale sharks elsewhere. While all sharks stayed in national waters, our results highlight the high mobility of juvenile whale sharks and demonstrate their connectivity across the Sulu and Bohol Seas, highlighting the importance of the area for this endangered species.

Satellite tagging highlights the importance of productive Mozambican coastal waters to the ecology and conservation of whale sharks.

The whale shark Rhincodon typus is an endangered, highly migratory species with a wide, albeit patchy, distribution through tropical oceans. Ten aerial survey flights along the southern Mozambican coast, conducted between 2004-2008, documented a relatively high density of whale sharks along a 200 km stretch of the Inhambane Province, with a pronounced hotspot adjacent to Praia do Tofo. To examine the residency and movement of whale sharks in coastal areas around Praia do Tofo, where they may be more susceptible to gill net entanglement, we tagged 15 juveniles with SPOT5 satellite tags and tracked them for 2-88 days (mean = 27 days) as they dispersed from this area. Sharks travelled between 10 and 2,737 km (mean = 738 km) at a mean horizontal speed of 28 ± 17.1 SD km day-1. While several individuals left shelf waters and travelled across international boundaries, most sharks stayed in Mozambican coastal waters over the tracking period. We tested for whale shark habitat preferences, using sea surface temperature, chlorophyll-a concentration and water depth as variables, by computing 100 random model tracks for each real shark based on their empirical movement characteristics. Whale sharks spent significantly more time in cooler, shallower water with higher chlorophyll-a concentrations than model sharks, suggesting that feeding in productive coastal waters is an important driver of their movements. To investigate what this coastal habitat choice means for their conservation in Mozambique, we mapped gill nets during two dedicated aerial surveys along the Inhambane coast and counted gill nets in 1,323 boat-based surveys near Praia do Tofo. Our results show that, while whale sharks are capable of long-distance oceanic movements, they can spend a disproportionate amount of time in specific areas, such as along the southern Mozambique coast. The increasing use of drifting gill nets in this coastal hotspot for whale sharks is likely to be a threat to regional populations of this iconic species.

Satellite tagging of rehabilitated green sea turtles Chelonia mydas from the United Arab Emirates, including the longest tracked journey for the species.

We collected movement data for eight rehabilitated and satellite-tagged green sea turtles Chelonia mydas released off the United Arab Emirates between 2005 and 2013. Rehabilitation periods ranged from 96 to 1353 days (mean = 437 ± 399 days). Seven of the eight tagged turtles survived after release; one turtle was killed by what is thought to be a post-release spear gun wound. The majority of turtles (63%) used shallow-water core habitats and established home ranges between Dubai and Abu Dhabi, the same area in which they had originally washed ashore prior to rescue. Four turtles made movements across international boundaries, highlighting that regional cooperation is necessary for the management of the species. One turtle swam from Fujairah to the Andaman Sea, a total distance of 8283 km, which is the longest published track of a green turtle. This study demonstrates that sea turtles can be successfully reintroduced into the wild after sustaining serious injury and undergoing prolonged periods of intense rehabilitation.

Some like it hot: Repeat migration and residency of whale sharks within an extreme natural environment.

The Arabian Gulf is the warmest sea in the world and is host to a globally significant population of the whale shark Rhincodon typus. To investigate regional whale shark behaviour and movements, 59 satellite-linked tags were deployed on whale sharks in the Al Shaheen area off Qatar from 2011-14. Four different models of tag were used throughout the study, each model able to collect differing data or quantities of data. Retention varied from one to 227 days. While all tagged sharks crossed international maritime boundaries, they typically stayed within the Arabian Gulf. Only nine sharks dispersed through the narrow Strait of Hormuz into the Gulf of Oman. Most sharks stayed close to known or suspected feeding aggregation sites over summer months, but dispersed throughout the Arabian Gulf in winter. Sharks rarely ventured into shallow areas (<40 m depth). A single, presumably pregnant female shark was the sole animal to disperse a long distance, crossing five international maritime boundaries in 37 days before the tag detached at a distance of approximately 2644 km from the tagging site, close to the Yemeni-Somali border. No clear space-use differentiation was evident between years, for sharks of different sizes, or between sexes. Whale sharks spent the most time (~66%) in temperatures of 24-30°C and in shallow waters <100 m depth (~60%). Sharks spent relatively more time in cooler (X2 = 121.692; p<0.05) and deeper (X2 = 46.402; p<0.05) water at night. Sharks rarely made dives deeper than 100 m, reflecting the bathymetric constraints of the Gulf environment. Kernel density analysis demonstrated that the tagging site at Al Shaheen was the regional hotspot for these sharks, and revealed a probable secondary aggregation site for whale sharks in nearby Saudi Arabian waters. Analysis of visual re-sightings data of tagged sharks revealed that 58% of tagged individuals were re-sighted back in Al Shaheen over the course of this study, with 40% recorded back at Al Shaheen in the year following their initial identification. Two sharks were confirmed to return to Al Shaheen in each of the five years of study.

Mobulid rays feed on euphausiids in the Bohol Sea.

Mobulid rays have a conservative life history and are caught in direct fisheries and as by-catch. Their subsequent vulnerability to overexploitation has recently been recognized, but fisheries management can be ineffective if it ignores habitat and prey preferences and other trophic interactions of the target species. Here, we assessed the feeding ecology of four mobulids (Manta birostris, Mobula tarapacana, M. japanica, M. thurstoni) in the Bohol Sea, Philippines, using stomach contents analysis of fisheries specimens landed between November and May in 2013-2015. We show that the mobulids feed heavily on euphausiid krill while they are in the area for approximately six months of the year. We found almost no trophic separation among the mobulid species, with Euphausia diomedeae as the major prey item for all species, recorded in 81 of 89 total stomachs (91%). Mobula japanica and M. thurstoni almost exclusively had this krill in their stomach, while M. tarapacana had a squid and fish, and Ma. birostris had myctophid fishes and copepods in their stomachs in addition to E. diomedeae. This krill was larger than prey for other planktivorous elasmobranchs elsewhere and contributed a mean of 61 364 kcal per stomach (±105 032 kcal s.e., range = 0-631 167 kcal). Our results show that vertically migrating mesopelagic species can be an important food resource for large filter feeders living in tropical seas with oligotrophic surface waters. Given the conservative life history of mobulid rays, the identification of common foraging grounds that overlap with fishing activity could be used to inform future fishing effort.

Population Structure, Abundance and Movement of Whale Sharks in the Arabian Gulf and the Gulf of Oman.

Data on the occurrence of whale sharks, Rhincodon typus, in the Arabian Gulf and Gulf of Oman were collected by dedicated boat surveys and via a public-sightings scheme during the period from 2011 to 2014. A total of 422 individual whale sharks were photo-identified from the Arabian Gulf and the northern Gulf of Oman during that period. The majority of sharks (81%, n = 341) were encountered at the Al Shaheen area of Qatar, 90 km off the coast, with the Musandam region of Oman a secondary area of interest. At Al Shaheen, there were significantly more male sharks (n = 171) than females (n = 78; X2 = 17.52, P < 0.05). Mean estimated total length (TL) for sharks was 6.90 m ± 1.24 (median = 7 m; n = 296). Males (7.25 m ± 1.34; median = 8 m, n = 171) were larger than females (6.44 m ±1.09; median = 7 m, n = 78; Mann-Whitney U test, p < 0.01). Of the male sharks assessed for maturity 63% were mature (n = 81), with 50% attaining maturity by 7.29 m and 100% by 9.00 m. Two female sharks of >9 m individuals were visually assessed as pregnant. Connectivity among sharks sighted in Qatari, Omani and UAE waters was confirmed by individual spot pattern matches. A total of 13 identified sharks were re-sighted at locations other than that at which they were first sighted, including movements into and out of the Arabian Gulf through the Strait of Hormuz. Maximum likelihood techniques were used to model an estimated combined population for the Arabian Gulf and Gulf of Oman of 2837 sharks ± 1243.91 S.E. (95% C.I. 1720-6295). The Al Shaheen aggregation is thus the first site described as being dominated by mature males while the free-swimming pregnant females are the first reported from the Indian Ocean.

Acoustic telemetry reveals cryptic residency of whale sharks.

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.

Laser photogrammetry improves size and demographic estimates for whale sharks.

Whale sharks Rhincodon typus are globally threatened, but a lack of biological and demographic information hampers an accurate assessment of their vulnerability to further decline or capacity to recover. We used laser photogrammetry at two aggregation sites to obtain more accurate size estimates of free-swimming whale sharks compared to visual estimates, allowing improved estimates of biological parameters. Individual whale sharks ranged from 432-917 cm total length (TL) (mean ± SD = 673 ± 118.8 cm, N = 122) in southern Mozambique and from 420-990 cm TL (mean ± SD = 641 ± 133 cm, N = 46) in Tanzania. By combining measurements of stranded individuals with photogrammetry measurements of free-swimming sharks, we calculated length at 50% maturity for males in Mozambique at 916 cm TL. Repeat measurements of individual whale sharks measured over periods from 347-1,068 days yielded implausible growth rates, suggesting that the growth increment over this period was not large enough to be detected using laser photogrammetry, and that the method is best applied to estimating growth rates over longer (decadal) time periods. The sex ratio of both populations was biased towards males (74% in Mozambique, 89% in Tanzania), the majority of which were immature (98% in Mozambique, 94% in Tanzania). The population structure for these two aggregations was similar to most other documented whale shark aggregations around the world. Information on small (<400 cm) whale sharks, mature individuals, and females in this region is lacking, but necessary to inform conservation initiatives for this globally threatened species.

Whale sharks target dense prey patches of sergestid shrimp off Tanzania.

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 (<2 mm) dominated background samples. The size structure was skewed towards larger animals representative of L.hanseni in feeding samples. Thus, whale sharks at Mafia Island target patches of dense, large, zooplankton dominated by sergestids. Large planktivores, such as whale sharks, which generally inhabit warm oligotrophic waters, aggregate in areas where they can feed on dense prey to obtain sufficient energy.