Many shark populations are in decline around the world, with severe ecological and economic consequences. Fisheries management and marine protected areas (MPAs) have both been heralded as solutions. However, the effectiveness of MPAs alone is questionable, particularly for globally threatened sharks and rays (‘elasmobranchs’), with little known about how fisheries management and MPAs interact to conserve these species. Here we use a dedicated global survey of coral reef elasmobranchs to assess 66 fully protected areas embedded within a range of fisheries management regimes across 36 countries. We show that conservation benefits were primarily for reef-associated sharks, which were twice as abundant in fully protected areas compared with areas open to fishing. Conservation benefits were greatest in large protected areas that incorporate distinct reefs. However, the same benefits were not evident for rays or wide-ranging sharks that are both economically and ecologically important while also threatened with extinction. We show that conservation benefits from fully protected areas are close to doubled when embedded within areas of effective fisheries management, highlighting the importance of a mixed management approach of both effective fisheries management and well-designed fully protected areas to conserve tropical elasmobranch assemblages globally. A survey of sharks and rays on coral reefs within 66 marine protected areas across 36 countries showcases that the conservation benefits of full MPA protection to sharks almost double when accompanied by effective fisheries management.

Overfishing remains a threat to coral reef fishes worldwide, with large carnivores often disproportionately vulnerable. Marine protected areas (MPAs) can restore fish populations and biodiversity, but their effect has been understudied in mesophotic coral ecosystems (MCEs), particularly in the Coral Triangle. Videos were analysed from baited remote underwater video systems deployed in 2016 to investigate the assemblage structure of large carnivorous fishes at shallow (4-12 m) and mesophotic (45-96 m) depths in two of the largest and most isolated MPAs in the Philippines: an uninhabited, fully no-take MPA enacted in 1988 (Tubbataha Reefs Natural Park) and an archipelagic municipality surrounded by an extensive but not fully no-take MPA declared in 2016 (Cagayancillo). Taxa focused on were groupers (Serranidae), snappers (Lutjanidae), emperors (Lethrinidae), jacks (Carangidae) and the endangered Cheilinus undulatus (Labridae). Mean abundance and species richness were not greater in TRNP than in Cagayancillo regardless of depth despite long-term protection in the former. Limited impacts of fishing in Cagayancillo may explain this result. Differentiation of fish assemblages was evident between TRNP and Cagayancillo but more obvious between depths at each location, probably due more to habitat than MPA effects. In Cagayancillo, overall carnivorous reef fish, grouper and jack mean abundance were 2, 2 and 10 times higher, respectively, at mesophotic depths, suggesting that MCEs can serve as deep refugia from fishing. These findings of differentiation between depths and higher abundance of certain taxa in mesophotic depths emphasize that MCEs are distinct from shallow reefs, serve as important habitat for species susceptible to overfishing and, thus, must be explicitly included in the design of MPAs. This study also highlights the value of maintaining strict protection of MPAs like TRNP for the Coral Triangle and an opportunity to safeguard intact fish assemblages in Cagayancillo by expanding its no-take zones.

A global survey of coral reefs reveals that overfishing is driving resident shark species toward extinction, causing diversity deficits in reef elasmobranch (shark and ray) assemblages. Our specieslevel analysis revealed global declines of 60 to 73% for five common resident reef shark species and that individual shark species were not detected at 34 to 47% of surveyed reefs. As reefs become more shark-depleted, rays begin to dominate assemblages. Shark-dominated assemblages persist in wealthy nations with strong governance and in highly protected areas, whereas poverty, weak governance, and a lack of shark management are associated with depauperate assemblages mainly composed of rays. Without action to address these diversity deficits, loss of ecological function and ecosystem services will increasingly affect human communities.

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.

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

The green turtle Chelonia mydas is an Endangered species that forms aggregations at neritic foraging sites where juveniles spend, in some cases, over a decade before moving to adult- or subadult-dominated foraging sites. Here, we used photographic identification, behavioural observations, paired-laser and stereo-photogrammetry to determine the population dynamics of a coastal aggregation of green turtles in Oslob, Philippines. We identified a total of 82 individual turtles between May 2012 and October 2018. We recorded behavioural observations during individual identification and found that turtles changed behaviour in 25% of instances. Some turtles were identified consistently throughout the study period, indicating strong site fidelity and residency. Modified maximum likelihood models suggest turtles spend the majority of their time within the study area (>92%), although some (8%) excursions outside the area do occur. Mark-recapture models suggest this foraging aggregation exhibits a pattern similar to that of a closed population. This is likely due to the extended residency periods of some individuals at the site, and the relatively low recruitment rate over time. We measured 18 turtles using paired-laser photogrammetry, with estimated straight carapace length of 55.3 cm, and mean growth rates of 3.4 cm yr(-1). Stereo-photogrammetry measurements were consistent and more accurate than visual or paired-laser photogrammetry estimates. Our results highlight the use of in-water methods to understand population dynamics of green turtles, and the importance of coastal, juvenile-dominated habitats for green turtles, as well as the need for effective conservation and management strategies to safeguard them.

Elasmobranchs are experiencing population declines worldwide because of anthropogenic stressors. Marine protected areas (MPAs) have been shown to benefit mesopredatory species of shark. The Tubbataha Reefs Natural Park (TRNP) is a United Nations Educational, Scientific and Cultural Organization World Heritage Site and the largest no-take MPA in the Philippines, yet information on the current status of elasmobranchs in the park is limited. Here, we investigate the abundance and biodiversity of elasmobranchs in TRNP using underwater visual census (UVC) and baited remote underwater video (BRUV) surveys. Twenty UVC were conducted in May 2015 (n = 10) and June 2016 (n = 10), documenting eight species of elasmobranchs with an average density of 7.18 sharks ha(-1) for both years. A total of 113 successful BRUV surveys were conducted in TRNP between March-June 2015 (n = 47) and April-June 2016 (n = 66). The BRUV surveys recorded 14 species with an average catch per unit effort of 2.09 elasmobranchs per hour for both years. Both survey methods report some of the highest densities of the grey reef shark Carcharhinus amblyrhynchos and the whitetip reef shark Triaenodon obesus documented worldwide. This study underlines the importance of large, well-managed MPAs such as TRNP for the conservation of reef-associated elasmobranchs. (C) 2018 National Science Museum of Korea (NSMK) and Korea National Arboretum (KNA), Publishing Services by Elsevier.

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.

In light of the global decline of mobulid populations and the necessity for sustainable fisheries management, baseline data for population dynamics were collected from a targeted fishery in the Bohol Sea, Philippines. This study focused on life-history parameters and reproductive cycles of four mobulid rays (Mobula thurstoni, M. japanica, M. tarapacana, and Manta birostris), and re-estimated their intrinsic population growth rates. Size and reproductive data were collected from 1,509 specimens (30% of catch) landed in two fishing seasons in 2015 and 2016. Size-at-birth was reviewed, and analysis of the embryos and follicles did not show any clear seasonality in the reproductive cycle, but supported an interbreeding interval. Females of all species matured at a larger size than males, and exhibited a larger size-at-pregnancy than -at-maturity. This delay in reproduction resulted in population growth rates lower than the actual rmax when based on size-at-pregnancy (rmat=rmax=0.016-0.055 year-1 and rpreg=0.008-0.044 year-1), and a population doubling time of 15.8-86.6 years. This study suggests that population growth rates previously reported were overestimated. In light of the Convention on International Trades of Endangered Species (CITES) and Convention on Migratory Species (CMS) assessments, while fisheries management should reflect the delayed maturation of these species and the slower population growth potential, at the current status of these population, the sustainability of any exploitation level seems unrealistic and strongly discouraged.

We report the occurrence of the rare and Endangered Aetomylaeus vespertilio (Bleeker, 1852) at the remote archipelago of Cagayancillo, Palawan in the Sulu Sea, Philippines. Only 1 previous  report of the species exists from the country, a juvenile specimen in the Pasil market, Cebu City in 2000. During an in-water shark and ray survey at Cagayancillo Island in April 2017, a female individual A. vespertilio with a disc width of approximately 200 cm, and a total length of approximately 400 cm was encountered along the reef’s edge at a depth of 12 m. This species wasconfirmed by its characteristic dorsal pattern of narrow stripes anteriorly and reticulated spots posteriorly. We confirm the occurrence of this rare and poorly understood species in the Sulu Sea and extend the current known range farther west than previously known.

Provisioning wildlife for tourism is a controversial yet widespread practice. We analysed the residency patterns of juvenile whale sharks (Rhincodon typus) in Oslob, Philippines, where provisioning has facilitated a large shark-watching operation since 2011. We identified 208 individual sharks over three years, with an average of 18.6 (s.d.=7.8, range=643) individuals sighted per week. Weekly shark abundance varied seasonally and peak-season abundance (approx. May-November) increased across years. Whale sharks displayed diverse individual site visitation patterns ranging from a single visit to sporadic visits, seasonal residency and year-round residency. Nine individuals became year-round residents, which represents a clear response to provisioning. The timing of the seasonal peak at Oslob did not align with known non-provisioned seasonal aggregations elsewhere in the Philippines, which could suggest that seasonal residents at Oslob exploit this food source when prey availability at alternative sites is low. Since prolonged residency equates to less time foraging naturally, provisioning could influence foraging success, alter distributions and lead to dependency in later life stages. Such impacts must be carefully weighed against the benefits of provisioning (i.e. tourism revenue in a remote community) to facilitate informed management decisions.

The whale shark, Rhincodon typus, is a charismatic umbrella species whose highly mobile nature is not yet fully understood. Whale sharks roam the Philippine archipelago with two major aggregations known to occur at Donsol and at a provisioning site in Cebu.This is the first description of a previously identified aggregation occurring off Panaon Island, Southern Leyte through the use of photographic identification. In total, 93 individual whale sharks were identified, with significant male bias (58%). The mean estimated total length of individuals was 5.72 1.02 m S.D., indicating a juvenile aggregation.Partial or complete fin amputations, potentially resulting from fishing lines, boat propellers or net entanglement, were observed on 27% of animals, highlighting some of the risks human activities can have on this threatened species. Multiple parallel scars, identified as propeller impact, were observed on 45% of animals.Dedicated research seasons in 2013 and 2014 yielded very different whale shark encounters with 366 in 2013 and 12 in 2014, yet highlighted the recurrence of individuals at the study site. Complemented by data collected through citizen science, maximum likelihood methods were used to model mean residency of whale sharks at Panaon Island of 27.04 days. The modelled lagged identification rate showed that many whale sharks return to the study site over time.Whale sharks from Panaon Island were identified through photo-ID and citizen science at other sites in the Philippines, as well as a match to Taiwan, representing the first international match through photo-ID in South-east Asia with a minimum distance covered of 1600 km.Given the highly mobile nature and recent exploitation of this species, management is recommended as a single unit regionally in South-east Asia. Additional research is needed to focus on the drivers of variation in encounters at whale shark aggregation sites. Copyright (c) 2016 John Wiley & Sons, Ltd.

Sample R code used to run permutation tests and PCoA to categorize whale shark residency patterns at the provisioning site. Data provided are three years of daily presence-absence data for Y1 ID frequent sharks (refer to manuscript)

Background Shark viewing tourism is a burgeoning global industry. Numerous operations provision sharks to increase sightings yet the effects of provisioning on shark behaviour are often poorly understood. At a unique provisioning site in Oslob, Cebu, Philippines, whale sharks (Rhincodon typus) have become the focus of a mass tourism operation since late 2011, receiving over 185,000 tourists a year. Approach To better understand and advise management bodies on the effects of provisioning on these animals, we used photographic identification (photo-ID) to assess the daily abundance and site visitation patterns of the sharks. Principal coordinate analysis (PCoA) based on co-occurrence matrices was used to investigate visitation patterns amongst individuals and define groupings. Modified maximum likelihood methods were then used to model mean residency for these different groups. Results Between March 31st 2012 and March 31st 2015, we identified 209 (165 males, 34 females and 10 of unknown sex) individuals at the site (a 0.065 km2 demarked area located 50 – 100 m from shore), with an average of 12.4 (SD = ± 4.8, range = 2 – 31) sharks sighted per day. Daily shark abundance increased over time and showed strong seasonal variation, with annual peaks occurring between May and November. We documented a spectrum of site visitation patterns ranging from single sighting (21% of sharks identified) to year-round residency (5% of sharks identified). For frequently sighted sharks, PCoA revealed two primary residency groupings: highly resident sharks (N = 9), which were seen year-round with few prolonged absences, and seasonal sharks (N = 21), which were primarily present between May and November. The daily abundance of highly resident sharks varied nonlinearly over time with no clear trend, while the daily abundance of seasonal sharks showed a clear increase during peak season in each year of the study. Mean residency for highly resident sharks was estimated at 55.5 ± 175.5 SE (95% CI 49.5 – 730.0) days, 58.3 ± 161.7 SE (95% CI 23.2 – 81.4) days for seasonal individuals, and 16.8 ± 1.8 SE (95% CI 14.8 – 22.2) days for infrequent sharks (N = 130). Conclusions The difference in residency patterns between frequently sighted and infrequent individuals was almost fourfold. This shift in residency could have long-term implications for ca. 15% of identified whale sharks at the site. We provide insight into the impacts of provisioning on shark movements and inform the debate surrounding the future of this controversial activity.

Determining the life history and population parameters of sea turtles typically involves the live capture of animals. Photo-identification (photo-ID) of free-ranging individuals is a cost-effective alternative to conventional tagging for mark-recapture studies that also minimises the risk of injury or stress to the animals. Paired laser-photogrammetry (photogrammetry) is a minimally invasive technique to size animals and has been proven effective in a range of different taxa. Photo-ID and photogrammetry were combined as minimally invasive techniques to determine life-history parameters of free-ranging green sea turtles Chelonia mydas at a site in Oslob, Cebu, Philippines. The pattern-recognition software I3S Pattern was used to validate results and confirm visual matching. A total of 38 individuals were identified at the site through photo-ID. Facial scutes remained unchanged for a min. of 3.6 years. Photogrammetry was employed to measure straight carapace lengths (SCLs) of a sample of 17 individuals, with a mean straight carapace length of 52.5 ± SD 1.1 cm. Modified maximum likelihood methods were used to model mean residency of 873 d at the site, with c. 12 individuals present at any one time. Based on photogrammetry measurements, individuals at the site were found to be of immature age class. This, along with the extended residency times, indicates that the site could be an important developmental habitat. Photo-ID is a valid minimally invasive technique for identifying individual C. mydas for mark-recapture analyses. The use of paired laser photogrammetry for categorising the life-history stage of an aggregation of sea turtles is demonstrated for the first time. Identification of such developmental habitats through the employment of these minimally invasive techniques is useful for conservation of the species.