
First described by J. Walbaum in 1792, manta rays are the largest living rays in the ocean. Measured by their wingspan, individuals may reach over 7m in disc width. Currently there are two recognized species of manta rays. Once thought to be a single worldwide species, intensive research into the matter revealed that at least two distinct species exist, Manta birostris and Manta alfredi. Manta birostris, which is commonly called the giant manta, is the larger and more widely distributed of the two species. Manta alfredi, or the reef manta, is the smaller species with a wingspan reaching only 5 meters. The species differ from one another in a number of ways including most noticeably dorsal and ventral colouration, texture of skin and morphology of denticles, number of teeth, and presence of a spine/stinger (albeit a non-functional one). However, major differences also exist in their behaviour, size at maturity, size of claspers (external sexual organs), and shape of their body (morphometrics). Detailed descriptions of the two recognized species and a comprehensive key to the genus are provided in our publication on the topic (Marshall et al. 2009). A third potential species (Manta sp. cf birostirs) is being examined in the Atlantic Ocean. Research into the validity of this third species is currently underway.

While manta rays have a circumglobal distribution (meaning that they are found across the globe) and are found in all oceans but the Arctic, they are mostly commonly found in tropical waters. The different species of manta rays have very different distributions, however, and are not commonly found residing together or using similar habitat.
Manta birostris appears to be more commonly found off shore and may be a more migratory species. This species has been documented to occur as far north as southern California and Rhode Island on the United States west and east coasts, Mutsu Bay, Aomori, Japan, the Sinai Peninsula, Egypt and the Azores Islands in the Northern Hemisphere and as far south as Peru, Uruguay, South Africa and New Zealand in the Southern Hemisphere.
Manta alfredi, is commonly sighted inshore, within a few kilometres of land. Found around coral and rocky reefs as well as along productive coastlines with consistent upwelling, tropical island groups, atolls and bays. This species is widespread in the Indian Ocean, with images and sightings of M. alfredi from the Red Sea in the north to Durban, South Africa in the south, and from mainland Thailand in the north to waters off Perth, Australia in the south. In the eastern and south Pacific, M. alfredi occurs from the Yaeyama islands, Japan in the north to the Solitary Islands, Australia in the south and is sighted as far east as French Polynesia south of the equator and the Hawaiian islands north of the equator. Two reports and photographs of M. alfredi from the north Atlantic off the Canary Islands and the Cape Verde Islands and historical reports and photos of M. alfredi off the coast of Senegal in north west Africa (Cadenat 1958) are the only evidence of populations of M. alfredi in Atlantic waters.
The two Manta species are sympatric in a few locations around the globe including Mozambique, Hawaii, and Indonesia but at most of these locations, geographical separation, fine-scale habitat use, or seasonal movement patterns typically keep them from coming in contact with one another. (See map)





Manta rays have very distinct body colouration including natural spot patterns on their ventral surface (stomach) that can be used to differentiate individual rays. Amazingly, aside from the natural colouration pattern most commonly seen in manta rays, two additional ‘colour morphs’ occur. Firstly there is a black or melanistic morph, where the entire ray is black aside from a variably sized blaze of white on its ventral surface. No other species of shark or ray has been observed to have a melanistic or black colour morph. While ‘black mantas’ are common in some regions, such as the indo pacific and along the west coast of the Americas, in some regions specimens are entirely absent, such as along the eastern coast of Africa where no black manta rays have been registered. Although far more rare than the melanistic morphs, a white or leucistic morph also occurs, where there is a notable absence of pigment on the body of the manta ray creating an overall whitish effect. These individuals are not true albinos as they still retain some pigment (particularly in their eyes) but they can be extremely devoid of pigment in some cases. It is not known when the genetic mutations that caused these rare colour morphs first occurred or why they have persisted over time, but it is certainly interesting that these colour morphs occur in both of the recognized species of Manta, suggesting that these genetic mutations occurred in an ancestral form (i.e. before the species split).
How old to manta rays live? At the moment, common techniques used to age other sharks and rays are not working for manta rays and it is unclear how long they live or at what age they reach sexual maturity. However, we expect that like many of the other large pelagic species of elasmobranchs, manta rays are long-lived and reach maturity at a late age. Researchers are busy trying to modify existing techniques and explore new ones so that we may finally figure out the longevity of both Manta species.
We do know that manta rays appear to spend all of their time in the water column; there is currently no evidence to suggest that they rest on the sea floor like other rays. They do appear to partition their time up doing different things. During the daylight hours, manta rays appear to most commonly spend their time inshore and are often seen at cleaning stations or feeding along the coast. During the evening, they appear to commonly move offshore or into deeper waters and it has been proposed that they may spend the majority of the nighttime hours feeding.
Manta rays, along with the devil rays, have the largest brain of any fish (which includes the other sharks and rays) relative to body size. While this might at first seem odd, it is becoming increasingly clear that these magnificent animals are highly social, a common trigger for rapid growth in brain size. Aside from the highly social and complex behaviours they exhibit, many of which we still do not fully understand, manta rays may also use sound to communicate to one another (intra-specific communication). There is evidence to suggest that the noise generated by breaching (jumping clear out of the water) may serve to signal to other rays in the vicinity.
It is often noted that manta rays look very different from other rays. The reason for their dramatic differences have a lot to do with their diet and their lifestyle, which ultimately required them to have a different body plan than their demersal relatives. As such, manta rays have undergone dramatic modification and as a result have many new and unusual adaptations. One of the most obvious differences lies in the diamond shape of their body; their massive pectoral fins are capable of propelling them through the water in the same fashion as a bird in the sky. These massive wings which have a mix of both white and red muscle (used for both quick bursts of speed and long sustained swimming) and provide manta rays with agility, maneuverability and speed and most importantly, enable them to undertake massive migrations. Manta rays are known to travel substantial distances sometimes over 200 kilometers a week and over 1,000 kilometers two months in a single direction. Manta rays also have what is known as lateral eye placement, meaning that instead of having eyes located on the top of their head (like demersal rays) they are located on the side of their head. This placement allows them greater range of vision in the pelagic environment where they spend their time. Despite this clever eye placement, manta rays do have distinct blind spots, which is evidenced by ‘rear approach’ predation injuries and line entanglement scars between their cephalic fins. Their cephalic fins themselves are a unique adaptation, only occurring in the mobulid rays. These lobes attached to the end of their head on either side of their mouth appear to aid in the capture of zooplankton, presumably making them more efficient feeders. Unlike other planktivorous elasmobranches like whale sharks and basking sharks, which are only capable of opening their mouths, manta rays are capable of angling these fins in different ways to direct water flow into their mouths. When not feeding, manta rays will roll these fins up to make themselves more hydrodynamically efficient whilst traveling. Early sailors and explorers thought that they resembled horns and thus named the group the ‘devil fish’, which they are still often referred to today. This often leads to negative feelings about these rays so we encourage using the name manta, which derives from the Spanish word for blanket or cloak.
Like many other large marine megafauna, manta rays are planktivores, feeding on small marine invertebrates called zooplankton and occasionally small fish. Manta rays employ many different types of feeding modes when trying to capture this tiny food source. Commonly, when plankton is distributed evenly in the water column they will simply feed at the surface or mid-water with their enormous mouths open sieving the tiny particulate animals from the water with specialized and highly modified gill rakers. These gill rakers prevent the food particles from passing over their gills (where oxygen is extracted from the water and ultimately exits via their ten gill slits). After being trapped, plankton particles are transported to the back of their mouth where they are swallowed. The opening of their esophagus or their ‘throat’ is approximately the size of an average human fist. It is for this reason that they cannot eat large items and pose no threat to humans. While the above-mentioned foraging method is the most commonly observed feeding mode, manta rays will alter their behaviour to suit conditions. If plankton is densely concentrated in one location manta rays are often observed barrel rolling in the water column (which facilitates them staying in one location or in a specific area). While making dramatic vertical loops, individuals are able to feed more efficiently on the concentrated plankton. Similarly, if plankton accumulates or is trapped just above the sea floor, manta rays are capable of feeding off the ground. Like a vacuum, the rays will almost scrape their huge terminal mouths along the seabed. They will also often use their cephalic fins in a unique manner, like a funnel, to direct plankton into their gaping mouths.
Despite swallowing their food whole, manta rays still possess teeth. Measuring only about 1-2mm in size, individual manta rays can have up to 4,000 of these minute teeth, which are embedded in a tooth band that stretches almost the entire length of the lower jaw. These teeth are not used for feeding but rather are used in their reproductive behaviour. Abrasive, like Velcro, male manta rays will grasp onto the pectoral fins of the female ray when mating using these teeth to help them maintain their grip. The teeth are not large or sharp enough to cut the female’s fin, but they do leave scratch marks that appear red when fresh (i.e when the female has been recently mated) and turn white to grey over time.

Underwater photography (Photo identification) is effective, efficient, and non-intrusive and is the underlying methodology behind most of our long-term studies. Each animal sighted on the local reefs is identified by a ‘fingerprint’ of markings on their ventral surface as well as by distinctive scars. Once individual manta rays have been identified, their details are logged in a computer database and their re-sightings tracked over time.
Over the last seven years, 700 individuals have been identified in southern Mozambique, making the Mozambican Manta Photo Database one of the largest and most comprehensive scientific manta database in the world. Over 40% of the total identified population has been re-sighted using photographic tagging techniques and acoustic technology, suggesting that there is a degree of philopatry or site fidelity to the immediate area or southern Mozambican region. At the studied aggregation site in southern Mozambique (off Inhambane) there is a strong sex bias in observed rays, with females out numbering males 3.5 to 1 on the reefs. Re-sighting data show that females are not only more prevalent in the area but were re-sighted more commonly than males. This stretch of coastline appears to be an adult aggregation site, with 90% of the male population and 55% of the identified female population being mature individuals.
Using photographic identification we have been able to conduct an in-depth study on the natural predation of manta rays by sharks at this location. In great contrast to other known aggregation sites, 76% of all identified mantas in this region bear significant injuries from shark attacks, the highest reported in the world. These attacks are made by a wide range of species including reefs sharks and larger predatory sharks like bull sharks and tiger sharks. For more information on this study please see our publication on this topic.
In conjunction with the bite mark analysis, observations and experiments on the cleaning behaviour and wound healing of manta rays has been conducted here in Mozambique. Cleaning stations (which are patches of reef frequented by manta rays for the purpose of being cleaned by small reef fish) are regularly monitored by our team and are also the best place for recreational scuba divers to have encounters with these giant rays. Dozens of cleaning stations have been identified in southern Mozambique, with many of these stations being regularly visited by manta rays. Over seven different species of cleaner fish clean manta rays along this coastline with each fish having a specific ‘job’. It has been found that these cleaner fish partition the manta ray’s body up as a common resource, with each species of fish cleaning a specific area and item off the manta ray’s body. Some of the identified individuals in the population have been noted to spend long periods of time cleaning. It is not uncommon for rays to spend several hours at cleaning stations per day allowing the cleaner fish to remove parasites and promote the healing of their bite wounds. Cleaning has been identified as a daytime activity with no rays recorded at the stations during the nighttime hours. For more information on our cleaning studies see our publication on the subject.
Long-term monitoring of manta rays at study sites in Mozambique has allowed us to ascertain population estimates along this stretch of coastline and track population composition seasonally. Using photo identification techniques manta rays are ‘visually tagged’ and ‘re-sighted’ over time, allowing us to examine the proportion of re-sighted individuals to new individuals that are continually being identified. These values can be used to estimate both the annual population size at our monitored aggregation sites and also the super population size (number of rays that may occur in this region).
Our current study is the first of its kind in Mozambique or even Africa, with our population estimates serving as the world’s first data on an identified manta population. To date, the population estimates for this immediate stretch of coastline suggest that we have approximately 800 reef manta rays (Manta alfredi) and approximately 600 giant mantas (Manta birostris) although giant manta rays are both observed less commonly and identified individuals re-sighted less often. To date, this is one of the largest populations of manta rays identified anywhere in the world. Combined with the fact that manta rays use this coastline year-round, southern Mozambique may offer divers and researchers alike the best opportunity to interact with these gentle giants.
While it is an extremely important first step to estimate the population sizes of manta rays, it is equally important to assess whether their numbers are stable, increasing or decreasing. This year we will be looking into a notable decline in the ‘sightability’ of rays over the last seven years along our famous ‘manta ray coastline’ and the factors that may be contributing to their gradual decline in these waters.

Again, applying photographic-identification methodology, many aspects of the reproductive ecology of reef manta rays, Manta alfredi, have been examined over the last half-decade. The region encompassing the study site was identified as a mating ground for M. alfredi back in 2003. Courtship and mating behaviour have been studied and described from many different events during the last seven consecutive summers. Over 70 different pregnant females have been sighted and 8 different newborns (1.3-1.6 metres) have been recorded on local reefs in the summer months. A detailed dissection of a full to term embryo was even conducted in late 2005 (see Marshall, A.D., Pierce, S.J. & Bennett, M.B. 2008).
The seasonality of mating events and the timing of parturition have been examined during this study. Based on observations of mating events and fresh mating scars on the pectoral fins of female rays, it is clear that Manta alfredi use the local habitat to breed and give birth. The distribution of these pectoral fin scars was highly biased and indicated a strong lateralised behavioural trait, with 99% of these scars occurring only on the left pectoral fin. No other elasmobranch has been reported to display behavioural lateralization (preference for a specific side- e.g. handedness in humans), making this both a surprising and fascinating discovery.
In southern Mozambique reef mantas give birth in the austral summer period after a gestation period of approximately one year. Reproductive periodicity in M. alfredi was most commonly found to be biennial, but a few individuals were seen to be pregnant in consecutive years, confirming an annual ovulatory cycle. The production of a single pup appears to be the normal situation for these rays, although two pups are conceived on occasion. Many aspects of the study have contributed to the limited baseline data currently available for this species (see Marshall, A. D., Compagno, L. J. V. & Bennett, M. B. 2009). The results of this research study on M. alfredi have highlighted the potential need for more conservative conservation strategies, with manta rays in general being one of the lest fecund elasmobranch species in the world.
Unfortunately, we have not been able to obtain meaningful information on the reproductive ecology of Manta birostris as only two pregnant females have been recorded during the study period. Clearly more information is needed on this giant manta ray, with next to nothing known about their biology or behaviour. Gathering data on this subject continues to be one of our primary objectives.
Despite their curious and tolerant nature, manta rays are elusive animals that are typically only encountered in surface waters where they are cleaning or feeding. As such we still know very little about these threatened species which continues to hamper our efforts to manage populations effectively, assess their threats and assign them an appropriate conservation status.
While most of our work is built around our non-intrusive photo identification program, various tagging programs have been started to establish how manta rays use near-shore environments. These programs also aim to identify critical habitats such as feeding areas, cleaning stations, and areas used for mating and birthing.
The first of our programs makes use of coded acoustic tags, which are placed on a random selection of individual rays. These tags relay information to listening stations (acoustic receivers) that we have strategically placed down on reefs or other important aggregation areas. Tagged animals will continually register (several times a minute) on these listening stations if they are using the area within 500 metres of one of the stations. Even if an individual only passes by, the stations will still record their presence. Over time, this acoustic-based system gives us an uninterrupted view of how these individuals use various habitats along the coastline, giving us an idea of which areas are heavily trafficked during which time periods, which area may serve as critical habitats that must be carefully monitored or protected, and which areas area used infrequently or seasonally. The more receivers used the better information we get back and it is our intention in 2010 to have over 20 listening stations deployed at aggregation sites along a 200 km stretch of coastline in southern Mozambique. This passive acoustic array will continue to gather valuable information on manta rays for years to come, with the batteries on tags lasting up to five years. Our local array will also link in with the Wildlife Conservation Society’s Indian Ocean ‘Meganet’ which has similar arrays operating in many locations in the western Indian Ocean, including Madagascar, Tanzania, Kenya and South Africa.
Over the long term this study will provide some of the data needed to support the design of a marine protected area in the region for manta rays and whale sharks. From a conservation angle, we will be able to target protection measures where they are most needed thus increasing the effectiveness of management efforts. By sharing movement data with other regional nodes of the meganet, we can look at the linkages between countries and start planning for more successful international management measures.
For the past half decade, our team has been collecting tissue samples of identified individuals in the Mozambican population. We are also collecting tissue samples from manta rays across the globe. To extract these samples we use highly efficient Hawaiian slings with biopsy tip that remove small plugs of tissue from the pectoral fins of individual rays. These samples allow for many types of studies. To date we have used our collection to examine the differences between the two populations of Manta alfredi and Manta birostris at our study site in Mozambique. We are also continuing our efforts to compare populations around the globe to better understand their movement patterns and relatedness. In the future we also hope to look in more detail at the relatedness of individuals at the population level here in Mozambique and overlay this information with the detailed ecological information that has been collected in the field.
Our feeding ecology team, team lead by Chris Rohner, will be trying to determine the dietary preferences and seasonal shifts in the diet of manta rays along this coastline. The ocean off Tofo is incredibly productive. As opposed to tropical seas, which are usually very clear, the waters along the Inhambane coastline are often full of plankton. Thanks to this increase productivity, we have a high abundance of both whale sharks and manta rays along the coastline. While manta rays and whale shark often feed together in the same area, observations have revealed that each of the animals have their own specialised strategies to make the most of the microscopic feast. Additionally, while preliminary observations suggest that both animals feed on the same or similar plankton species, it is still unclear if each of the species have dietary preferences.
Scientists from the Foundation began in 2009 to look into the plankton dynamics in our local waters. Even though this is work is still in progress, a nice success story of a little planktonic animal called Diacavolinia longirostris is emerging. This tiny gastropod is often seen by divers on their safety stop in the area and our researchers have long suspected that the mantas might particularly seek out and feed on this zooplankton species. Lately, our team has been building more and more evidence to support is theory. Diacavolinia is often seen when large feeding aggregations of manta rays occur and we have observed individual manta rays ingesting them. Plankton tows in the immediate vicinity of barell-rolling mantas have further supported this theory as has our analysis of manta waste (yes…we even collect this in the field! Is there no end to our dedication?).
Comparative manta ray research has been conducted in other international destinations including, Mexico, Australia, Yap, Brazil, the Maldives and Indonesia. Currently we have many collaborative studies underway, with many others due to begin in 2010, including those which will take our team to southern Brazil, Ecuador, Australia, Egypt, Mexico, Thailand, and the United States.
Some of this research is directed at comparing population sizes and structures, while other research focuses more on obtaining valuable biological data from mantas killed in target fisheries. Our largest worldwide collaboration currently underway is aimed at learning more about the movement patterns and migratory behaviour of the elusive giant manta rays at key seasonal aggregation sites. With generous funding from the Save Our Seas Foundation our research team will be using satellite telemetry to tag and track giant manta rays as they roam the world’s oceans. It is our hope that this study will ultimately shed light on their habitat use, diving abilities and their long range swimming abilities. This is the first study of its kind and the collaborative nature of the project will allow for many important comparisons as well as make it a robust first look at this mysterious ocean wanderer.
As human pressures increase worldwide manta ray populations have declined in many areas where they were once common. There is a notable contrast between the few reasonably protected populations, such as in Yap, the Maldives and the Hawaiian Islands, where economically valuable ecotourism and dive operations exist, and populations in areas where fisheries target species, such as along the coast of Africa, India, South East Asia, and Central America. Sadly, due to many of their life history traits mantas are highly vulnerable to over-fishing, and there looms a genuine threat of localised extinction of certain populations.
Currently there are no comprehensive management programs for manta rays anywhere in the world, yet they are listed by the IUCN as ‘near threatened’ to extinction. This conservation assessment has not been updated since the discovery of two distinct species. The recent reclassification of the genus has major implications for the conservation assessment of the two species. Each species faces different and specific threats in various regions of the world, and the worldwide IUCN status of the genus requires urgent revaluation in light of this revision. Our team is currently leading efforts to make this reassessment.
Acquiring accurate information on population dynamics, lifespan, reproductive parameters, growth rates and natural mortality rates is crucial to understanding the conservation requirements of a species. Additional information on the life history of manta rays is sorely needed to supplement the paucity of existing data, particularly for giant mantas (Manta birostris) for which little biological information is known. Site fidelity, movement patterns and habitat usage are also essential pieces of information needed to properly manage populations, once the biological parameters of a species are known. This type of information can highlight an animal’s potential susceptibility to fishing pressures and help determine critical habitats and seasonal migration routes. Elucidating these types of large behavioral trends in elusive pelagic animals can prove challenging but with the help of modern technology we are better equipped to try than at any other point in history.
At the end of the day, management itself is the true challenge and real effort needs to be made to protect populations or critical habitats once they have been identified. In doing this, the animals themselves need to be protected from intense anthropogenic pressure of any kind. While directed fishing pressure is the most obvious threat, management must also extend to recreational boating and fishing, as well as, activities involving human interactions (e.g. scuba diving). All such activities must be regulated in order to ensure that even cryptic threats are monitored and minimized, the health and behaviour of animals is not dramatically altered and the sustainability of eco-tourism is maintained.
Eco-tourism has proven to be an excellent way of generating a positive, sustainable balance between protecting populations of animals and creating an economically viable form of tourism. In 2002, the Big Island of Hawaii was reported to have generated over $2.5 million US dollars in combined revenues from manta related activities in their coastal waters. Surveys in the Maldives have reported that the local economy generates approximately $7.8 million US dollars in combined revenue from manta ray related activities in their islands. These figure are astonishing when compared to the $250-500 US that a dead manta can yield in the fish markets across the world. Some of the directed fisheries that supply this market can harvest as many as 1,500 individuals per annum. Manta rays are predominantly used for their skin, cartilage, and gill rakers, which are used in bogus Chinese medicinal products and for ornamental goods. In some areas, like along the eastern coast of Africa, manta rays are also still taken in subsistence fisheries.
Manta ray populations are frequently small in size and they are one of the least fecund species of sharks or rays (low number of offspring). Intense fishing pressure can threaten the stability of local populations and their numbers can be decimated in the span of a few years. Our conservation efforts in Mozambique aim to raise the profile of manta rays in the region and ultimately formally protect both species of manta rays in Mozambican waters. Long-term we hope to assist with the development of marine protected areas for manta rays and whale sharks in critical habitats such as the Tofo coastline south of Inhambane. Globally, foundation scientists will continue to contribute through our involvement with the IUCN redlist assessment for manta rays. We also will continue to collaborate with international manta ray programs to uncover population trends, assess threats, and develop management plans.
To get more involved we suggest the following: