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Future research should include a comprehensive survey of the West African coastline to collect specimens and investigate diversity within the region.

For the purposes of this paper, we identify existing research and knowledge gaps for M. In addition to the currently recognized M. Conclusive molecular support and clear morphological descriptions based on sufficient population and genomic sampling are still lacking Hinojosa-Alvarez et al. However, a formal description of M. Rediscovery of cryptic diversity within manta rays in the early Twenty-first Century was the result of detailed morphological observation of many individuals of several populations combined with genetic analyses of multiple genes Marshall et al.

The lack of equivalent studies for the remaining devil rays indicates that currently undetected cryptic diversity within the group cannot be ruled out, and thus we recommend additional detailed taxonomic studies for mobulid rays generally. Further, population level sampling enables opportunities for studies to define Evolutionarily Significant Units ESUs for conservation prioritization. To support future morphometric and meristic-based studies, we strongly encourage research groups and collections to improve reference material at the population level to effectively catalog mobulid diversity.

Life-history information is needed to estimate extinction risk, maximum population growth rates in data-poor species, and for performing stock assessments Musick, ; Pardo et al. These slow-growing rays have very low rates of reproduction and long maturation times Notarbartolo di Sciara, ; Marshall and Bennett, ; Dulvy et al. Key life history parameters such as age at maturity, growth rate, lifespan, mortality both natural and fisheries-induced and fecundity are lacking for most mobulid species Table 1.

Estimating these parameters is crucial for assessing the vulnerability and demography of exploited populations Campana et al. Wherever possible, life-history parameter estimates should be obtained and applied to management at the population level, as biological characteristics are likely to vary both among species and locations with variable environmental conditions. Regional differences may be especially relevant in broadly distributed species such as M. Available estimates of mobulid extinction risk suggest that mobulid ray populations are unlikely to withstand current levels of fishing mortality, even in small-scale artisanal fisheries Dulvy et al.

Therefore, accurate estimates of life history parameters are urgently needed to assess the effectiveness of potential fisheries management scenarios.

In elasmobranchs, growth is usually modeled by estimating the age of individuals of known size disc width in rays by counting growth bands on vertebral centra Figure 2 or other hard parts such as fin spines Cailliet et al.

The vertebral centra of devil rays tend to be poorly calcified, which has hindered the use of this technique so far. Currently, only one data set has provided size-at-age estimates to model growth rates using vertebrae from the caudal region of M.

Further efforts should assess this method in other species and locations, since the extent of calcification may vary. Researchers should strive to obtain samples from a broad size-range and estimate growth parameters using a multi-model approach i.

In data-poor situations or when a size-class is poorly represented, back-calculation, or Bayesian methods may allow for adequate estimates Cailliet et al. An important assumption is that each pair of opaque and translucent bands is deposited annually Cailliet et al. Validating this assumption in mobulid rays could be attempted by injection of a fluorochrome label e. Alternatively, radiocarbon 14 C dating of archival devil ray hard parts could provide a method of age validation by tracing this dated chemical marker released in high concentrations during atomic bomb testing in the 60—70 s Campana et al.

Additionally, researchers could evaluate the feasibility of using cesium isotopes released during the Fukushima nuclear accident to assist with age validation for mobulids in the western Pacific Neville et al. Until validation is achieved, researchers can test band-pair deposition consistency across years as a means of verification Cailliet et al. Figure 2. Age and growth in mobulid rays.

See section 2. Aging elasmobranchs using vertebrae relies on access to fished specimens and therefore is not appropriate in many wild mobulid ray populations. Direct estimates of growth rate can be obtained in-situ from morphometric measurements of resighted individuals over known time increments, based on visual size estimates Kitchen-Wheeler et al. These mark-recapture data with associated size estimates can be used to fit growth curves e. Therefore, further efforts should aim to obtain accurate size estimates over longer time frames to estimate growth rates.

It will also be important to obtain accurate estimates of size-at-birth from neonates or late-term embryos across populations since these are necessary to fit two-parametric growth models, which are preferable to three-parametric models that sometimes provide unrealistic estimates of size at birth in elasmobranchs Smart et al. Mortality is an essential parameter to understand population dynamics, and it is important for demographic models and stock assessments to separate natural mortality from fishing mortality.

There are a number of methods researchers can use to estimate mortality in mobulid rays depending on both data availability and type. In data-rich wild populations, mark-recapture via photo identification and acoustic telemetry can be used to estimate survival and natural mortality of specific life stages e. In fisheries scenarios, catch curve analysis of landed specimens allows for direct estimates of total mortality natural plus fishing mortality e.

Many indirect methods exist to estimate mortality based on other life history parameters Pauly, ; Kenchington, The most relevant approaches to mobulid research likely include estimation of natural mortality from maximum age e. The life history information highlighted in this section is important for developing management strategies that prioritize life stages for conservation, and for testing the ability of proposed conservation approaches to enhance population viability. Matrix models are a common tool for population viability analysis, and can be particularly useful in data-poor scenarios that are common among mobulid rays Heppell et al.

As key life history data become available, future research on mobulids should use sensitivity, elasticity, and perturbation analyses to identify life stages with significant contributions to overall population viability in order to direct management actions Heppell, ; Frisk et al. Additionally, researchers could assess the impact of specific management approaches by quantitatively evaluating the increase in the growth rate of a population given reduced mortality of specific life stages e.

Further, in cases where time-series of abundance or relative abundance are not available which is the case for many mobulid populations , these life history parameters can be used to calculate the maximum growth rate of a population Pardo et al.

This can be compared with total mortality to determine population trajectories and extinction risk Pardo et al. The reproductive biology of mobulid rays is known mostly from the reef manta ray M.

Courtship and copulation in reef manta rays are common around cleaning stations and reef habitats Marshall and Bennett, ; Deakos et al. Observations of courtship and mating remain rare in other mobulid species Yano et al. A gestation period of 1 year has been observed in both wild and captive M. All mobulids are thought to give birth to a single large pup, with rare observations of twins Marshall et al. Parturition is often followed immediately by copulation in captive and wild M.

In wild populations of M. The fecundities of mobulids other than M. Size segregation appears to be common across mobulid species, with juveniles absent from most field studies or present at different times or locations than adults Notarbartolo di Sciara, ; Deakos, , with the exception of M.

In terms of movements, habitat use and general life history data, the juvenile life stage of all mobulid species remains poorly studied. Maturity assessments of mobulids are based primarily on external morphological cues, such as changes in clasper morphology in males, and the presence of visual signs of mating or pregnancy in females Deakos, ; Marshall and Bennett, ; Stevens, Figure 3.

However, physiological sexual maturity may not coincide with the external signs of sexual maturation. Physiological markers such as changes in steroid levels have been successfully adopted as indicators of reproductive status in elasmobranchs Sulikowski et al. Ultrasonography has also been widely adopted to define gonadal maturity in a variety of fish species Bryan et al.

Future research addressing mobulid reproduction and fecundity could consider applying these methods, particularly in species where individual identification to track reproductive status through time is not possible. If used on a sufficiently large scale, these approaches could help estimate the proportion of females within a population contributing to annual recruitment and, for data collected over extended periods of time, may provide information on the timing and seasonality of the reproductive cycle Whittamore et al.

In combination with photogrammetry, ultrasonography and endocrinology would enable more reliable estimates of age and size at maturity. Figure 3. Mobulid reproduction. Mating, pupping and nursery grounds are not well defined for mobulids. See section 3. In several mobulid species, reproductive activity peaks seasonally and may occur more often at social aggregation sites such a seamounts or cleaning stations Yano et al. However, significant knowledge gaps remain in our understanding of mobulid courtship and mating behavior.

Long-term monitoring of areas where opportunistic courtship activity has been documented previously may reveal more seasonal mating grounds for certain species, and targeted surveillance of social aggregation sites would be a natural starting place to undertake these studies.

Little is known about parturition in mobulid rays. To date, no natural wild birth for any mobulid species has been observed. Advanced technology may be required to determine if specific pupping grounds exist, or if females give birth in any area with suitable conditions. Researchers could use satellite and acoustic tags to evaluate the movements and habitat use of near-term females and reveal behavioral changes preceding pupping events.

Additionally, animal-borne video camera systems e. In many locations, juvenile M. Deakos, G. Stevens unpublished and occasionally in offshore habitats R.

Rubin unpublished segregated from adult populations. Additionally, juvenile M. To date two locations have been identified that meet established criteria for elasmobranch nursery habitats Heupel et al. Future work should target locations where juvenile mobulids are known to aggregate or are frequently present, and determine whether these sites meet the criteria for formal classification as a nursery habitat as defined by Heupel et al. Given the apparent affinity for protected coastal and lagoon habitats in several species, aerial drone surveys could be an efficient and cost-effective method for identifying candidate sites.

The identification of mating and pupping grounds may also aid in the identification of juvenile habitats and nursery grounds. For example, large annual aggregations of M.

Juvenile and potential young-of-year individuals have been observed along the coast adjacent to these aggregations Notarbartolo di Sciara, G. Stewart, pers. If mating pupping, and nursery habitats are identified, these critical habitats should be protected to prevent changes in natural behavior, to ensure that pathways to these areas are not obstructed and to ensure the safety of individuals within these areas.

With increasing concern for the general status of global mobulid populations, estimates of abundance, and breeding stock along with monitoring of long-term population trends are crucial measures required to develop effective management and conservation strategies. Current population information for manta rays is predominantly gathered through diver counts and photo-identification photo-ID surveys of targeted populations at predictable aggregation sites. However, photo-ID is limited to species with individually unique markings e.

Therefore, there is a need to investigate additional methods to assess population trends, especially for devil rays. Figure 4. Photo identification of mobulids. Oceanic manta rays M. Both species have chevron a,c and black b morphs.

See section 4. Of all mobulid species, M. The large variation in body pigmentation patterns, presumed to be unique to each individual from birth, facilitates the cataloging of individuals and the creation of photo-ID databases Kumli and Rubin, ; Kitchen-Wheeler et al. The popularity of in-water interactions with mobulids has also resulted in a high incidence of citizen science contributions to regional photo-ID databases from recreational divers Kumli and Rubin, ; Couturier et al.

In addition to benefiting researchers by expanding data collection spatially and temporally without additional cost or resource use, citizen science promotes public awareness, and engagement in conservation actions Bonney et al. The opportunistic manner by which these sightings are reported, however, can impede the use of mark-recapture analyses that require precise sampling designs to control for effort and information on absence as well as presence of individuals Cooch and White, Researchers should carefully consider the analytical methods they plan to employ and the key parameters they wish to recover, and plan their data collection accordingly rather than relying purely on opportunistic sightings.

Furthermore, a key limiting factor to mark-recapture analysis is recapture rate: the probability of observing an individual animal on subsequent events. Patchy data often result in low recapture rates, and this may lead to non-estimable parameters or low precision around estimates, precluding the determination of trends. The use of auxiliary data within mark-recapture frameworks can help to overcome limitations with low recapture rates.

For example, the incorporation of acoustic tagging data into mark-recapture analysis of tagged seven-gill sharks Notorhynchus cepedianus facilitated abundance estimation for this species Dudgeon et al. Several mobulid research programs already include tagging studies see Movements section and future research programs could consider integrating these study designs to enable best use of data across multiple studies.

Rigorous sampling designs can also increase the utility of citizen science data. For example, focusing citizen science efforts during particular time periods can increase data input into analysis frameworks, as well as investigating presence-only models to include opportunistic sightings Mengersen et al.

Ongoing surveys of well-studied populations should be continued to enable long-term trends to be assessed. For many mobulid species, assessing population trends through individual identification photo-ID, tag, and release is impossible due to extremely low recapture probabilities resulting from infrequent encounters, diver avoidance behaviors, or lack of distinct physical markings. Other approaches therefore need to be considered.

In populations of mobulids that may exhibit high site affinity and recapture rates, researchers could evaluate the efficacy of external tags or implanted PIT tags Chapman et al. Indirect or relative estimates of abundance can be generated through: a count surveys SPUE: sighting per unit effort of live animals e. As life history parameters and population estimates become available for mobulids, researchers should use traditional stock assessment methods to evaluate the status and trends of populations Methot and Wetzel, However, this will most likely only be possible in fisheries that have high observer coverage or bycatch reporting e.

In more common data-poor scenarios, researchers could take advantage of existing length- and age-based assessment methods such as catch curve analysis Pardo et al. Where robust estimates of current population sizes are available e. New developments in genomic approaches show promise for facilitating population size estimates for mobulids. DNA can be extracted from small amounts of tissue collected from live or dead animals as well as preserved components such as gill plates. High throughput sequencing enables thousands of genetic markers to be sequenced most commonly focusing on single nucleotide polymorphisms—SNPs , which can then be used in multiple population genetic analyses.

One approach is estimating genetic effective population size Ne , which provides the number of breeding individuals to be used as a metric of population viability. Strong concordance has been demonstrated between ecological and genetic estimates of abundance for some elasmobranch species Portnoy et al.

Investigations are ongoing for M. Armstrong unpublished and Mozambique S. Venables unpublished. The advantage of Ne is that only a single population sample is required to generate a point estimate. Theoretically, changes in Ne with temporal sampling can demonstrate population trends e. However, given that demography, ecology, and reproductive mode can all influence Ne estimates Wang, , future research is needed to investigate the sensitivity of Ne to these biological features in mobulids and provide guidelines on minimum sampling requirements.

Another emerging molecular approach to estimating population size is close-kin mark-recapture CKMR; Bravington et al. Genomic data are used to identify kin pairs including parent-offspring, full siblings and half-siblings. CKMR has been applied to natural populations of bluefin tuna Bravington et al. As with Ne , CKMR only provides information about adults and requires sampling a substantial proportion of the population to identify kinship pairs and generate abundance estimates Bravington et al.

However, combining CKMR methods with auxiliary data, such as acoustic telemetry data, can provide estimates of key demographic parameters needed to estimate total population size Hillary et al. With few published examples for CKMR, sampling guidelines are not yet available, however several elasmobranch studies are underway Ovenden et al. The greatest contributors to direct mortality of mobulid rays across their range are targeted and bycatch fisheries Croll et al. At least 13 fisheries in 12 countries specifically target mobulids, and at least 30 fisheries in 23 countries capture mobulids as bycatch Hall and Roman, ; Croll et al.

They are targeted or retained for the Asian medicine trade as well as consumption of meat Croll et al. The attraction of mobulids to productive tropical and subtropical habitats where target species such as tunas aggregate, along with their distribution in the epipelagic zone, make them vulnerable to fisheries capture Croll et al. Mobulids are targeted or caught as bycatch in virtually every fishing gear type, including small-scale fisheries characterized by the use of driftnets, gillnets, harpoons, gaffs, traps, trawls, and longlines; and large-scale fisheries using driftnets, trawls and purse seines Croll et al.

The level of bycatch depends greatly on the fishing method used, with the highest bycatch rates reported from gillnets and purse seiners Croll et al. The slow population growth rates of mobulid populations largely exclude them from considerations of sustainable targeted fisheries Dulvy et al. Consequently, future research should focus on assessing bycatch risks and mitigation efforts as well as the impacts of existing fisheries on mobulid populations. Figure 5. Mobulid fisheries.

Mobulid rays are captured incidentally in virtually all fishing gear types, and are also targeted for their meat and gill plates in some countries. See section 5. While fisheries regulations have sought to prevent the retention and landing of mobulid rays, the vast majority of mobulid captures are a result of unintentional bycatch Croll et al.

Physical and biological traits of rays i. In the past, onboard handling practices have resulted in high levels of post-release mortality Hall and Roman, ; Poisson et al. Further work to identify and implement less harmful handling practices could prevent a significant amount of onboard mobulid mortality. Evaluating survival rates of mobulids released alive in non-target fisheries will help guide management decisions, as simply banning landings will not be an effective strategy in cases where a high proportion of individuals die after release.

Initial studies of M. Researchers should evaluate the post-release mortality of mobulids captured incidentally in a variety of gear types, as fishing gear such as gill nets with long soak times may have considerably higher post-release mortality rates than long lines or purse seines. Additionally, future studies should evaluate the impact of handling and release methods and relevant environmental and operational covariates on mobulid post-release mortality in order to develop release guidelines that maximize survival.

Post-release mortality can be evaluated using pop-off satellite tags Francis and Jones, or blood chemistry analyses Hutchinson et al. In coastal fisheries where released mobulids are more likely to remain in range of acoustic receivers, researchers could also consider the use of acoustic tags to evaluate survivorship rates Skomal, Accelerometer tags may also be useful in the quantification of post-release behavior and mortality and the impacts of handling methods in cases where instrument recovery is feasible Whitney et al.

As relationships between directly observable covariates e. Identifying areas of overlap between mobulid hotspots and fisheries could help reduce mobulid bycatch rates. Concurrent satellite tracking of focal species and vessel monitoring systems can provide insights into bycatch risk and identify key locations for mitigation and management action Queiroz et al.

Onboard fisheries observers provide far more data-rich scenarios in which to assess bycatch risk, especially in commercial fisheries with high observer coverage such as the IATTC tuna purse seine fleet Hall and Roman, ; Croll et al.

These observer records allow for detailed species distribution models that can reveal relationships between species abundance and key environmental variables, both for bycatch and target species Scales et al. Future studies should identify the regions with both the highest overall mobulid bycatch rates, and the highest mobulid bycatch to target catch ratios. This information will allow managers to develop spatio-temporal and ideally dynamic management approaches with the greatest ecological and conservation value for mobulids and the lowest economic loss from reduced catches of target species.

However, to take full advantage of fisheries observer data to develop species-specific habitat and distribution models, data collection protocols need to be standardized and species identification training for observers emphasized.

Researchers could work with regional fisheries management organizations RFMOs to develop a comprehensive, standardized data collection manual for mobulids that ensures all relevant variables, including release methods, are collected and can be compared among regions and fisheries. Additionally, RFMOs should be encouraged to implement detailed mobulid identification training courses, and researchers could follow up with molecular identification of captured specimens to determine identification error rates.

Unfortunately, these approaches will not cover mobulid mortality onboard smaller tuna purse seine vessels that don’t carry observers, or for fisheries targeting species outside the purview of the current RFMOs e. Many of these vessels operate in nearshore productive waters where mobulids aggregate, and are likely to represent substantial unreported mobulid bycatch.

Further research is necessary to describe the nature and quantity of this bycatch e. Bycatch mitigation methods have not been adequately explored for mobulids, and proposals for preventing interactions between fishing gear and mobulids through technological innovations or gear modifications are needed. Strong associations with the thermocline in M. However, the feasibility of limiting gear depth in commercial fisheries is questionable, and therefore alternate proposals should be developed and tested for mobulids more generally.

Bycatch mitigation has been developed for many non-target marine species such as seabirds in long-line fisheries e. For elasmobranchs more generally, there have been various trials, such as the apparently unsuccessful use of rare-earth metals to deter sharks from baited hooks Jordan et al. Some studies suggest that the use of light-emitting diodes in or near the ultraviolet range may reduce bycatch of elasmobranchs in gill net fisheries Jordan et al. Although in-depth research into population genetic structure is lacking for mobulids, the limited studies of genetic connectivity indicate that populations are spatially structured Stewart et al.

Consequently, fisheries management of mobulid rays may be more effective and relevant at the stock level, rather than simply along political or geographic boundaries Reiss et al. More genetic studies are needed to identify genetically distinct populations of mobulid rays to support regional management strategies. As traditional gene sequencing methods may not reveal population structure in mobulids Kashiwagi et al.

Given that many of the smaller mobulid species are challenging to find and sample in typical field expeditions, fisheries may provide the best opportunities to obtaining such sample representation. For example, tissue, or tail samples collected by observers in tuna fisheries could provide the extensive geographic coverage and the large sample sizes necessary for investigating genetic connectivity.

Additionally, there is strong evidence for a positive correlation between genetic diversity and a population’s resilience to extinction Frankham, ; von der Heyden, and overexploited fish species have been shown to have lower genetic diversity than closely related species that are not overharvested Pinsky and Palumbi, We therefore encourage researchers to evaluate genetic diversity across a gradient of fishing pressure to establish if and where fishing has led to a loss of diversity and population bottlenecks e.

This will help prioritize vulnerable populations for conservation and management action. Studies of general patterns of effective population sizes and descriptions of the demographic histories of mobulids can establish valuable baseline reference points.

Understanding the spatial dynamics and habitat use patterns of animal populations is key to successful conservation and management Cooke, ; Ogburn et al. This theme has been a dominant focus of mobulid ray research over the past decade, with 17 published studies documenting movements of individuals using a combination of photographic mark-recapture methods and electronic tracking.

Using acoustic and satellite telemetry, movements of 1, individual rays of 6 species have been investigated in 10 distinct regions throughout their range Figure 6. Together, these studies highlight the large movement capacity of manta and devil rays and their use of broad geographic ranges including coastal and pelagic waters Croll et al. High rates of site residency and fidelity have been demonstrated, particularly in manta rays Jaine et al.

The drivers underlying these large seasonal aggregations, common to both manta and devil rays, remain elusive. Identifying common physical and biological processes that underlie movements and aggregative behavior across species will allow for improved identification and characterization of critical habitats.

Studies of fine-scale habitat use at aggregation sites have revealed that individuals visit these areas to attend shallow cleaning stations, engage in courtship behavior, and forage on ephemeral food resources Dewar et al. Figure 6. Mobulid telemetry effort to date. The mobulid tagging effort information was sourced from published literature on the topic as well as from other unpublished mobulid tracking studies known to the authors and their collaborators summarized in Table S1.

See section 6. Technological advances in satellite tagging Musyl et al. Most studies of mobulid movements have employed pop-up satellite archival transmitting PSAT tags Table S1 , which typically yield long retention times but coarse position estimates i.

Using these, mobulid rays have been successfully tracked for periods of up to a year, revealing regional philopatric movements and strong affinity to shelf edge habitats in manta rays Braun et al. In contrast, towed Argos-linked tags require long tethers to facilitate surface satellite communications, leading to shorter deployments poor tag retention but higher resolution position estimates whenever the tag surfaces. These have been successfully used to investigate short-term i.

More recently, towed tags enabled with fast-acquisition GPS technology e. Erdmann pers. Fastloc positions require less surface time, allowing for shorter tethers and longer retention times than Argos-linked tags, but do not transmit archived high-resolution GPS positions until the tag detaches from the animal.

Recent improvements to geolocation and subsequent state space models are quantifying and reducing the levels of uncertainty associated with light-based geolocation Patterson et al. It is likely that with the increasing capacity to make fine-scale observations, movement patterns will indicate significant overlap with anthropogenic threats or sensitive areas Braun et al.

Researchers could therefore incorporate metrics such as the Human Impact Index Halpern et al. Fine-scale habitat use assessments will be facilitated by increased use of Argos-linked and Fastloc tags, and researchers could make efforts to improve retention times for these tag types, perhaps by implementing new attachment methods such as dorsal fin mounts e.

In addition to contributing important spatial information about the conservation needs of these species, such information will enhance our understanding of the global connectivity of mobulid populations with respect to gene flow and potential isolation.

Compared with satellite tags, the affordability and longer battery life of acoustic tags enables the collection of larger datasets on mobulid movements, site fidelity and habitat use than studies employing satellite tags. The increased battery life of acoustic transmitters up to 10 years will also greatly benefit efforts to monitor site fidelity and residency in mobulid species that are rarely encountered and are often only tagged opportunistically e.

However, the long-term attachment of external tags on mobulid rays over months to years remains a challenge. Future studies should consider internal tagging where possible e. Kessel et al. Additionally, acoustic tags only provide positions when they are in range of acoustic receiver stations, limiting their utility for wide-ranging mobulids that routinely travel outside of receiver arrays. Researchers could make use of collaborative regional acoustic sensor networks to expand monitoring capacity e.

Large scale collaborative efforts will inform the design of management strategies on national or smaller scales Lea et al. The ability of researchers to identify and predict areas of critical habitat for mobulid rays will be improved with increasing amounts of movement data collected through telemetry studies Hays et al. Density maps e. Historical observations of mobulid presence and aggregation events may provide novel and temporal insight into species distributions.

Researchers should consider Unmanned Aerial Vehicle UAV technology or aerial surveys to provide additional opportunities to collect quantitative and behavioral data of mobulids Hodgson et al. It is highly likely that oceanographic features and their influence on the distribution of patchy prey resources play a key role in driving the behavior of planktivores Sims et al. Studies should include oceanographic and biological information, such as prey field density and frontal systems, in their spatial analyses to expand our current understanding of mobulid movement patterns e.

Together, these data can be used in ecosystem niche modeling to identify potential new regions of importance to mobulid species Panigada et al. The position uncertainty inherent in light-based geolocation tags impedes the incorporation and comparison of oceanographic data with movements as prey fields and oceanographic features can vary widely within the error radius of a single position estimate. Increased use of Argos-linked transmitters with more precise position estimates may facilitate the inclusion of these important physical and biological features into spatial analyses.

It is crucial to gain a broader understanding of mobulid habitat use to better understand the effects of anthropogenic activities Graham et al. Ontogenetic shifts in movements and habitat use are common in elasmobranchs Grubbs, Juvenile mobulids are rarely seen and presumably exhibit size or age segregation until they approach maturity Stewart et al. Consequently, critical habitat use and movements likely vary among life stages in mobulid rays. Future studies could seek to address these ontogenetic differences, in particular by filling knowledge gaps in the movements and habitat use of juvenile and young-of-year mobulids.

Ongoing efforts to monitor mobulid movements through photo-ID Kumli and Rubin, ; Couturier et al. Fortunately, strong collaborative momentum within the scientific community and developments in analytical tools are now allowing for telemetry studies to take place at unprecedented scales, tracking movements of a multitude of marine species across oceans and over several years e. Acoustic and satellite telemetry studies continue to provide insights into the movement and spatial use patterns of mobulids Croll et al.

This site and species specificity in combination with limited sample sizes does not sufficiently support our understanding of broad-scale patterns at the population or species level. We recommend that future studies emphasize multi-species and multi-region tagging efforts to define critical habitats, resolve patterns in movement and connectivity among populations, improve our understanding of species-specific drivers of movement, and support efforts for management and bycatch reduction.

Determining drivers of mobulid aggregative and migratory behavior and how they are linked to foraging opportunities can help identify where these species are most susceptible to direct or incidental capture in fisheries Rohner et al. Mobulids are restricted to tropical and subtropical oceans, which in many regions are oligotrophic and may have lower zooplankton biomass than temperate and polar systems Moriarty and O’Brien, This means that many populations of mobulids must find high biomass food patches in a dilute food environment.

Stomach content analysis suggests that mobulids are opportunistic feeders that exploit zooplankton and, sometimes, teleost prey that are available in high densities Figure 7. For example, in Mexico and the Philippines, several mobulid species M.

Similarly, studies evaluating the prey community alongside feeding mobulids showed that M. Results from studies using stable isotopes and fatty acids as trophic tracers support direct observation data that mobulids are predominantly consumers of zooplankton, but that fishes make up a notable portion of their diet in some regions Sampson et al.

Further, some studies have suggested that M. Diving capacities and behaviors of several mobulid species also support the hypothesis that these animals can exploit prey in deep oceanic layers Canese et al. To help predict spatiotemporal distributions of mobulids, future studies should aim to identify common characteristics among their prey species such as community composition, biomass, size spectra and swimming capacity.

Further, examination of diving capacities of mobulids will provide a better understanding of different foraging strategies across their range and how these are affected by environmental variability e. Knowledge of prey landscapes alongside drivers and types of movement is key to habitat modeling efforts and the implementation of effective conservation measures for mobulids.

Figure 7. Foraging behavior of mobulid rays. The majority of direct observations of foraging are from epipelagic waters. However, a broad range of methods including isotope and fatty acid analysis, archival satellite tagging, stomach content analysis and submersible observations indicate that mesopelagic prey are important diet items for mobulids. References for this figure are listed in Table S2.

See section 7. Understanding oceanographic mechanisms that influence the distribution of mobulids will be important for identifying critical habitats that warrant special protection see section 6. Movements: Critical habitats. However, the temporal lag between primary productivity, which can be observed via satellite, and secondary productivity i.

Incorporating prey e. When possible, researchers should strive to collect prey data concurrently with mobulid sightings and behavioral data McCauley et al. However, in studies using telemetry and fisheries data this may not be feasible. Instead, researchers could take advantage of existing oceanographic datasets with high-resolution prey data e.

We encourage researchers to consider what prey data are available at or near their proposed study region, and even to consider specifically targeting regions with large oceanographic datasets for future mobulid research. Additionally, we encourage researchers to collaborate with oceanographers and cruise planners to help ensure that large-scale zooplankton data collection efforts are most relevant to studies of higher trophic level organisms including mobulids.

Bridging the gap between oceanographic data and mobulid movements and foraging behavior will allow for more robust analyses of distribution, critical habitat use, bycatch risk, and climate impacts.

Given the limitations of temporal coverage in stomach content analyses, a number of studies have employed stable isotope analysis and fatty acid profiling to examine the diet of mobulid rays Sampson et al. While diet-tissue discrimination values and turnover rates have been examined for several elasmobranchs Hussey et al. Captive feeding experiments on mobulids under controlled conditions in aquaria are needed to evaluate turnover times, diet tissue discrimination factors, and validate the use of other tissue types that can be sampled by non-invasive means for biochemical analyses, such as mucus Burgess et al.

Standardized protocols for sample preservation e. Future biochemical work on mobulid species could also consider compound-specific isotope analysis, a relatively new analytic tool in dietary studies Mcmahon et al. This technique provides a finer resolution on the origin of individual components e.

In all isotopic studies, adequate sampling of the food web and candidate diet items identified in observational studies and stomach contents is necessary to parameterize mixing models and quantify dietary contributions Burgess et al.

By combining several biochemical techniques and using large datasets, future studies will bring new understanding on trophic markers resulting from the different analyses and allow for more robust data interpretations on the dynamics of energy transfer in the pelagic environment.

Over the past decade, there has been an increased effort in using tagging technology to assess the vertical movement of mobulids and associated mesopelagic foraging ability.

Telemetry studies of other marine megafauna species have identified that the deep scattering layer Goldbogen et al. For mobulid species, all telemetry studies have shown that individuals spend most of their time within the upper 50 m of the water column where the likelihood of encountering fishing gear is greatest.

Several studies suggest that mesopelagic food sources comprise a major part of mobulid diet Burgess et al. Additionally, vertical migrators such as euphausiids and myctophids that appear to be an important prey source for mobulids may allow them to feed on mesopelagic prey sources without accessing mesopelagic environments Burgess et al. Many manta ray aggregation sites are adjacent to shelf breaks, and telemetry data suggest that individuals may depart coastal habitats at night Burgess, ; Couturier et al.

Obtaining new information on mesopelagic prey sources may require targeted field expeditions that sample prey at depth, which can be done with opening-closing nets, or could employ technology such as submersibles Stewart et al. A better understanding of mobulids’ reliance on these food sources and habitats will lead to improved characterization of aggregation sites, as well as habitat and species distribution models. As technology develops, so too does the ability to track and monitor mobulids in remote and deep realms of the ocean.

Animal-borne cameras Stewart et al. In studies of the diving behavior of other marine species, accelerometers have been successfully used to remotely assess diving energetics in the whale shark Gleiss et al. We therefore encourage researchers to apply accelerometers to future studies of mobulid feeding and diving behavior. This will require advances in attachment methods that allow accelerometer orientation to remain stable on mobulids, and exploring options for longer-term deployments and instrument recovery.

New sensory systems can also be incorporated into existing telemetry devices. For example, fluorometers embedded into satellite tags can provide in situ measurements of phytoplankton fluorescence that can be used to calculate chlorophyll- a concentrations and to assess primary productivity levels alongside animal movement data Lander et al.

Mobulid rays, as filter feeders, may be susceptible to ingesting marine pollutants and contaminants such as persistent organic pollutants POPs , heavy metals, and microplastics. Pollutants enter the marine environment through wastewater, poor industry practices, and degradation of marine debris, among other sources. Microplastics contain added pollutants and toxins such as phthalates, bisphenol A, flame retardants, styrenes, and adsorb and concentrate persistent organic pollutants POPs and heavy metals from the marine environment Worm et al.

Many dissolved pollutants are lipophilic, persistent in the environment, and have the capacity to bio accumulate up the food chain, with large bodied, long-lived species at high risk to exposure Niimi, Mobulid habitats, including foraging grounds, overlap with microplastic pollution hotspots in many cases Germanov et al. However, rates of plastic ingestion by mobulids, bioaccumulation of pollutants, and the impacts of plastic pollution on mobulid biology, ecology, and population viability have not been studied.

Future research should work toward identifying the scale of these problems and quantifying impacts so they can be evaluated in a management context. In locations with high densities of floating microplastics, mobulids may directly ingest microplastics Figure 8.

Additionally, zooplankton can be contaminated with pollutants and toxins Fossi et al. This suggests that mobulids may be secondary consumers of microplastics and associated pollutants even if they are foraging in locations or at depths that do not have high densities of floating microplastics.

Future research should seek to quantify ingestion rates of microplastics in mobulids. This could be accomplished by evaluating microplastic densities and contamination in prey assemblages by sampling prey alongside feeding mobulids.

Additionally, measuring the abundance of microplastic particles in stomach contents of mobulids landed in fisheries could provide estimates of direct ingestion.

In order to evaluate whether ingestion leads to bioaccumulation in tissues, researchers should measure phthalate, heavy metal, and POP levels in mobulid tissue biopsies from wild populations and samples collected from fisheries. Previous studies found elevated levels of some heavy metals in mobulid tissues Essumang, , ; Ooi et al. Future research in mobulids could evaluate levels of phthalates, POPs and heavy metals in mobulid tissues across gradients of microplastic pollution to determine how bioaccumulation scales with pollutant levels in the environment and to identify heavily impacted populations.

Sources and aggregating mechanisms of plastic pollution vary between coastal and offshore environments, leading to differing levels of overlap between microplastic hotspots and mobulid habitats Germanov et al. Researchers could investigate differences in pollutant contamination between coastal mobulids e. Once trophic interactions between mobulids and pollutants are better understood, researchers could endeavor to identify high-risk areas for mobulids following existing approaches for other taxa such as sea turtles Schuyler et al.

Figure 8. Sub-lethal impacts on mobulids. Entanglement in fishing gear can cause substantial damage and scarring a , including amputations of cephalic fins and damage to eyes b. The impacts of these stressors on mobulid foraging success, reproductive output, and population viability are poorly studied.

See sections 8 and 9. To face this new challenge I surrounded myself with experts and professionals in this field. We found an innovative solution. Ocean became my passion and I made it my profession as sailor. All world seas are now in danger. We have to face this new challenge of fighting ocean pollution”. Email: contact theseacleaners.

Serge Menard suggests that 40 of these giant tankers might clean up the oceans. This is a larger version of Yvan Bourgon’s Manta sea cleaners, save that the smaller ship is solar, sail and wind turbine powered. This website is provided on a free basis as a public information service. COFL is a charity without share capital.

 
 

MANTA SEA CLEANERS YVAN BOURGNON PLASTIC FACTORY SHIP

 
Methot, R. However, we hope that this identification of high priority knowledge gaps will stimulate and focus future mobulid research.

 

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