Manta ray

Manta rays are large rays belonging to the genus Mobula (formerly
its own genus Manta). The larger species, M. birostris, reaches 7 m Manta ray
(23 ft) in width, while the smaller, M. alfredi, reaches 5.5 m (18 ft). Temporal range: 23–0 Ma [1]
Both have triangular pectoral fins, horn-shaped cephalic fins and
PreꞒ Ꞓ O S D C P T J K PgN
large, forward-facing mouths. They are classified among the
Early Miocene to present
Myliobatiformes (stingrays and relatives) and are placed in the
family Myliobatidae (eagle rays).

Mantas are found in warm temperate, subtropical and tropical

waters. Both species are pelagic; M. birostris migrates across open
oceans, singly or in groups, while M. alfredi tends to be resident and
coastal. They are filter feeders and eat large quantities of
zooplankton, which they gather with their open mouths as they
swim. However, research suggests that the majority of their diet
(73%) actually comes from mesopelagic sources. Gestation lasts Manta alfredi at Dharavandhoo,
over a year and mantas give birth to live pups. Mantas may visit Maldives
cleaning stations for the removal of parasites. Like whales, they
breach for unknown reasons.

Both species are listed as vulnerable by the International Union for

Conservation of Nature. Anthropogenic threats include pollution,
entanglement in fishing nets, and direct harvesting for their gill
rakers for use in Chinese medicine. Their slow reproductive rate
exacerbates these threats. They are protected in international waters
by the Convention on Migratory Species of Wild Animals, but are
more vulnerable closer to shore. Areas where mantas congregate are
popular with tourists. Only a few public aquariums are large enough Manta birostris at Hin Daeng,
to house them. Thailand

Conservation status

Contents
Taxonomy and etymology
Species Vulnerable (IUCN 3.1)[3]
Fossil record
Scientific classification
Biology
Kingdom: Animalia
Appearance and anatomy
Lifecycle Phylum: Chordata
Behavior and ecology Class: Chondrichthyes
Distribution and habitat Order: Myliobatiformes
Conservation issues Family: Mobulidae
Threats
Status Genus: Manta
Relation with humans Bancroft, 1829 [2]
Aquariums
Type species
Tourism
Cephalopterus manta
See also
Bancroft, 1829
References
Species
External links

Manta alfredi
Taxonomy and etymology Manta birostris
†Manta hynei
The name "manta" is
Hexatrygon Portuguese and Spanish
for mantle (cloak or
blanket), a type of
Plesiobatis
blanket-shaped trap
traditionally used to catch
Urobatis rays.[5] Mantas are
known as "devilfish" Range of manta rays
because of their horn-
Pteroplatytrygon shaped cephalic fins, which are imagined to give them an "evil"
appearance.[6]
Potamotrygon Manta rays are members of the order Myliobatiformes which
consists of stingrays and their relatives.[4] The genus Manta is part
Elipesurus of the eagle ray family Myliobatidae, where it is grouped in the
subfamily Mobulinae along with the Mobula devil rays.[7] In 2017,
an analysis of DNA, and to a lesser degree, morphology, found that
Dasyatis Mobula was paraphyletic with respect to the manta rays, and they
recommended treating Manta as a junior synonym of Mobula.[8]
Gymnura Mantas evolved from bottom-dwelling stingrays, eventually
developing more wing-like pectoral fins.[9] M. birostris still has a
Aetoplatea vestigial remnant of a sting barb in the form of a caudal spine.[10]
The mouths of most rays lie on the underside of the head, while in
mantas, they are right at the front.[11] Manta rays and devil rays are
Myliobatis the only ray species that have evolved into filter feeders.[4]

Aetobatus Species

The scientific naming of mantas has had a convoluted history,

Aetomylaeus
during which several names were used for both the genus
(Ceratoptera, Brachioptilon Daemomanta, and Diabolicthys) and
Rhinoptera species (such as vampyrus, americana, johnii, and hamiltoni). All
were eventually treated as synonyms of the single species Manta
birostris.[12][13][14] The genus name Manta was first published in
Mobula 1829 by Dr Edward Nathaniel Bancroft of Jamaica.[12] The specific
name birostris is ascribed to Johann Julius Walbaum (1792) by some
Manta
Phylogeny of stingrays authorities and to Johann August Donndorff (1798) by others.[14]
(Myliobatiformes)[4] The specific name alfredi was first used by Australian zoologist
Gerard Krefft, who named the manta after Prince Alfred.[13][15]

Authorities were still not in agreement and some argued that the black
color morph was a different species from the mostly white morph.
This proposal was discounted by a 2001 study of the mitochondrial
DNA of both.[16] A 2009 study analyzed the differences in
morphology, including color, meristic variation, spine, dermal
denticles (tooth-like scales), and teeth of different populations. Two
distinct species emerged: the smaller M. alfredi found in the Indo-
Pacific and tropical east Atlantic, and the larger M. birostris found
throughout tropical, subtropical and warm temperate oceans.[10] The
former is more coastal,[3] while the latter is more ocean-going and Manta alfredi with mouth closed,
migratory.[17] A 2010 study on mantas around Japan confirmed the cephalic fins rolled and ventral
morphological and genetic differences between M. birostris and M. surface showing distinctive markings
alfredi.[18]

A third possible species, preliminarily called Manta sp. cf. birostris, reaches at least 6 m (20 ft) in width, and
inhabits the tropical west Atlantic, including the Caribbean. M. birostris and it occurs in sympatry.[10] More
recently, DNA studies published in 2018 suggest that the genus Manta should be nested within Mobula, and
that six existing species of Mobula should be consolidated into three.[19]

Fossil record

While some small teeth have been found, few fossilized skeletons of manta rays have been discovered. Their
cartilaginous skeletons do not preserve well, as they lack the calcification of the bony fish. Only three
sedimentary beds bearing manta ray fossils are known, one from the Oligocene in South Carolina and two
from the Miocene and Pliocene in North Carolina.[1] Remains of an extinct species have been found in the
Chandler Bridge Formation of South Carolina. These were originally described as Manta fragilis, but were
later reclassified as Paramobula fragilis.[20]

Biology

Appearance and anatomy

Manta rays have broad heads, triangular pectoral fins, and horn-
shaped cephalic fins located on either side of their mouths.[13] They
have horizontally flattened bodies with eyes on the sides of their
heads behind the cephalic fins, and gill slits on their ventral
surfaces.[13][21] Their tails lack skeletal support and are shorter than
their disc-like bodies.[21] The dorsal fins are small and at the base of
the tail. The largest mantas can reach 1,350 kg (2,980 lb).[13] In both
species, the width is about 2.2 times the length of the body; M.
Side view of M. birostris
birostris reaches at least 7 m (23 ft) in width, while M. alfredi reaches
about 5.5 m (18 ft).[22] Dorsally, mantas are typically black or dark in
color with pale markings on their "shoulders". Ventrally, they are
usually white or pale with distinctive dark markings by which individual mantas can be recognized.[10] Their
skin is covered in mucus which protects it from infection.[23]:2 All-black color morphs are known to exist.[21]
A pink manta ray has been observed in Australia's Great Barrier Reef and scientists believe this could be due
to a genetic mutation.[24] The fish, spotted near Lady Elliot Island, is the world's only known pink manta
ray.[25][26]

The two species of manta differ in color patterns, dermal denticles,

and dentition. M. birostris has more angular shoulder markings, larger
ventral dark spots on the abdominal region, charcoal-coloured ventral
outlines on the pectoral fins, and a dark colored mouth. The shoulder
markings of M. alfredi are more rounded, while its ventral spots are
located near the posterior end and between the gill slits, and the mouth
is white or pale colored. The denticles have multiple cusps and
overlap in M. birostris, while those of M. alfredi are evenly spaced
and lack cusps. Both species have small, square-shaped teeth on the
M.alfredi with cephalic fins rolled up
lower jaw, but M. birostris also has enlarged teeth on the upper jaw. (Yap, Micronesia)
Unlike M. alfredi, M. birostris has a caudal spine near its dorsal
fin.[10]

Mantas move through the water by the wing-like movements of their pectoral fins, which drive water
backwards. Their large mouths are rectangular, and face forward as opposed to other ray and skate species
with downward-facing mouths. The spiracles typical of rays are vestigial, and mantas must swim continuously
to keep oxygenated water passing over their gills.[23]:2–3 The cephalic fins are usually spiraled but flatten
during foraging. The fish's gill arches have pallets of pinkish-brown spongy tissue that collect food
particles.[13] Mantas track down prey using visual and olfactory senses.[27] They have one of the highest
brain-to-body mass ratios[28] and the largest brain size of all fish.[29] Their brains have retia mirabilia which
may serve to keep them warm.[30] M. alfredi has been shown to dive to depths over 400 metres (1,300 ft),[31]
while their relative Mobula tarapacana, which has a similar structure, dives to nearly 2,000 metres
(6,600 ft);[32] the retia mirabilia probably serve to prevent their brains from being chilled during such dives
into colder subsurface waters.[33]

Lifecycle

Mating takes place at different times of the year in different parts of

the manta's range. Courtship is difficult to observe in this fast-
swimming fish, although mating "trains" with multiple individuals
swimming closely behind each other are sometimes seen in shallow
water. The mating sequence may be triggered by a full moon and
seems to be initiated by a male following closely behind a female
while she travels at around 10 km/h (6.2 mph). He makes repeated
efforts to grasp her pectoral fin with his mouth, which may take 20 to
30 minutes. Once he has a tight grip, he turns upside-down and
M. alfredi group in the Maldives
presses his ventral side against hers. He then inserts one of his
claspers into her cloaca, where it remains for 60–90 seconds.[34] The
clasper forms a tube which channels sperm from the genital papilla; a
siphon propels the seminal fluid into the oviduct.[35] The male continues to grip the female's pectoral fin with
his teeth for a further few minutes as both continue to swim, often followed by up to 20 other males. The pair
then parts.[34] For some reason, the male almost always grasps the left pectoral fin, and females often have
scars that illustrate this.[23]:8–9
The fertilized eggs develop within the female's oviduct. At first, they are enclosed in an egg case while the
developing embryos absorb the yolk. After hatching, the pups remain in the oviduct and receive additional
nutrition from milky secretions.[36] With no umbilical cord or placenta, the unborn pup relies on buccal
pumping to obtain oxygen.[37] Brood size is usually one or occasionally two. The gestation period is thought
to be 12–13 months. When fully developed, the pup resembles a miniature adult and is expelled from the
oviduct with no further parental care. In wild populations, an interval of two years between births may be
normal, but a few individuals become pregnant in consecutive years, demonstrating an annual ovulatory
cycle.[36] The Okinawa Churaumi Aquarium has had some success in breeding M. alfredi, with one female
giving birth in three successive years. In one of these pregnancies, the gestation period was 372 days and at
birth the pup had a width of 192 cm (76 in) and weight of 70 kg (150 lb).[38] In southern Africa, M. birostris
males mature at 4 m (13 ft) while females reach maturity slightly over that.[39]:57 In Indonesia, M. birostris
males appear to mature at 3.75 m (12 ft), while female mature around 4 m (13 ft).[40] In southern Africa, M.
alfredi matures at widths of 3 m (10 ft) for males and 3.9 m (13 ft) for females.[39]:42 In the Maldives, males of
M. alfredi mature at a width of 2.5 m (8 ft 2 in), while females mature at 3 m (9.8 ft).[3] In Hawaii, M. alfredi
matures at a width of 2.8 m (9 ft 2 in) for males and 3.4 m (11 ft) for females.[41] Female mantas appear to
mature at 8–10 years.[3][17] Manta rays may live as long as 50 years.[22]

Behavior and ecology

Swimming behavior in mantas differs across habitats: when travelling

over deep water, they swim at a constant rate in a straight line, while
further inshore, they usually bask or swim idly around. Mantas may
travel alone or in groups up to 50. They may associate with other fish
species, as well as sea birds and marine mammals.[21] Mantas
sometimes breach, leaping partially or entirely out of the water.
Individuals in a group may make aerial jumps one after the other.
These leaps occur in three forms – forward leaps where the fish lands M. alfredi foraging with mouth
head first, similar jumps with a tail first re-entry, or somersaults.[13] opened wide and cephalic fins
The reason for breaching is not known; possible explanations include spread
mating rituals, birthing, communication, or the removal of parasites
and commensal remoras (suckerfish).[23]:15

Manta rays are filter feeders as well as macropredators. On the

surface, they consume large quantities of zooplankton in the form of
shrimp, krill, and planktonic crabs. In deeper depths, mantas consume
small to medium-sized fish.[42] When foraging, it slowly swims
around its prey, herding it into a tight "ball", and then speeds through
the bunched organisms with a wide-open mouth.[21] If a ball is
particularly dense, a manta may somersault through it.[23]:13 While
M. alfredi at a coral reef cleaning feeding, mantas flatten their cephalic fins to channel food into their
station with fish picking off parasites mouths and the small particles are collected by the tissue between the
gill arches. As many as 50 individual fish may gather at a single,
plankton-rich feeding site.[13] Tests have shown that around 27
percent of the diet of M. birostris is from the surface while around 73 percent is at deeper depths.[42] Mantas
are themselves preyed upon by large sharks and by killer whales. They may also be bitten by cookiecutter
sharks,[23]:17 and harbor parasitic copepods.[23]:14

Mantas visit cleaning stations on coral reefs for the removal of external parasites. The ray adopts a near-
stationary position close to the coral surface for several minutes while the cleaner fish consume the attached
organisms. Such visits most frequently occur when the tide is high.[43] In Hawaii, wrasses provide the
cleaning; some species feed around the manta's mouth and gill slits, while others address the rest of the body
surface.[23] In Mozambique, sergeant major fish clean the mouth, while butterflyfishes concentrate on bite
wounds.[39]:160 M. alfredi visits cleaning stations more often than M. birostris.[39]:233 Individual mantas may
revisit the same cleaning station or feeding area repeatedly[44] and appear to have cognitive maps of their
environment.[27]

In 2016, scientists published a study in which manta rays were shown to exhibit behavior associated with self-
awareness. In a modified mirror test, the individuals engaged in contingency checking and unusual self-
directed behavior.[45]

Distribution and habitat

Mantas are found in tropical and subtropical waters in all the world's major oceans, and also venture into
temperate seas. The furthest from the equator they have been recorded is North Carolina in the United States
(31°N) and the North Island of New Zealand (36°S). They prefer water temperatures above 68 °F (20 °C)[21]
and M. alfredi is predominantly found in tropical areas.[10] Both species are pelagic. M. birostris lives mostly
in the open ocean, travelling with the currents and migrating to areas where upwellings of nutrient-rich water
increase prey concentrations.[46]

Fish that have been fitted with radio transmitters have traveled as far as 1,000 km (620 mi) from where they
were caught, and descended to depths of at least 1,000 m (3,300 ft).[47] M. alfredi is a more resident and
coastal species. Seasonal migrations do occur, but they are shorter than those of M. birostris.[3] Mantas are
common around coasts from spring to fall, but travel further offshore during the winter. They keep close to the
surface and in shallow water in daytime, while at night they swim at greater depths.[21]

Conservation issues

Threats

The greatest threat to manta rays is overfishing. M. birostris is not evenly distributed over the oceans, but is
concentrated in areas that provide the food resources it requires, while M. alfredi is even more localized. Their
distributions are thus fragmented, with little evidence of intermingling of subpopulations. Because of their long
lifespans and low reproductive rate, overfishing can severely reduce local populations with little likelihood that
individuals from elsewhere will replace them.[17]

Both commercial and artisanal fisheries have targeted mantas for their meat and products. They are typically
caught with nets, trawls, and harpoons.[17] Mantas were once captured by fisheries in California and Australia
for their liver oil and skin; the latter were used as abrasives.[13] Their flesh is edible and is consumed in some
countries, but is unattractive compared to other fish.[48] Demand for their gill rakers, the cartilaginous
structures protecting the gills, has recently entered Chinese medicine.[49] To fill the growing demand in Asia
for gill rakers, targeted fisheries have developed in the Philippines, Indonesia, Mozambique, Madagascar,
India, Pakistan, Sri Lanka, Brazil, and Tanzania.[48] Each year, thousands of manta rays, primarily M.
birostris, are caught and killed purely for their gill rakers. A fisheries study in Sri Lanka and India estimated
that over 1000 were being sold in the country's fish markets each year.[50] By comparison, M. birostris
populations at most of the key aggregation sites around the world are estimated to have significantly fewer
than 1000 individuals.[51] Targeted fisheries for manta rays in the Gulf of California, the west coast of Mexico,
India, Sri Lanka, Indonesia, and the Philippines have reduced populations in these areas dramatically.[17]

Manta rays are subject to other anthropogenic threats. Because mantas must swim constantly to flush oxygen-
rich water over their gills, they are vulnerable to entanglement and subsequent suffocation. Mantas cannot
swim backwards, and because of their protruding cephalic fins, are prone to entanglement in fishing lines, nets,
ghost nets, and even loose mooring lines. When snared, mantas often attempt to free themselves by
somersaulting, tangling themselves further. Loose, trailing line can wrap around and cut its way into its flesh,
resulting in irreversible injury. Similarly, mantas become entangled in gill nets designed for smaller fish.[52]
Some mantas are injured by collision with boats, especially in areas where they congregate and are easily
observed. Other threats or factors that may affect manta numbers are climate change, tourism, pollution from
oil spills, and the ingestion of microplastics.[17]

Status

In 2011, mantas became strictly protected in international waters

because of their inclusion in the Convention on Migratory Species of
Wild Animals. The CMS is an international treaty organization
concerned with conserving migratory species and habitats on a global
scale. Although individual nations were already protecting manta
rays, the fish often migrate through unregulated waters, putting them
at increased risk from overfishing.[53] The IUCN declared M. birostris
to be 'vulnerable with an elevated risk of extinction' in November
2011.[54]
M. birostris at Hin Daeng, near Phi
Phi Islands, Thailand
In the same year, M. alfredi was also classified as vulnerable with
local populations of fewer than 1000 individuals and little or no
interchange between subpopulations.[3] The Manta Trust is a UK-
based charity dedicated to research and conservation efforts for manta rays. The organization's website is also
an information resource for manta conservation and biology.[55]

Besides these international initiatives, some countries are taking their own actions. New Zealand has banned
the taking of manta rays since the introduction of the Wildlife Act in 1953. In June 1995, the Maldives banned
the export of all ray species and their body parts, effectively putting a stop to manta fishing, as there had not
previously been a fishery for local consumption. The government reinforced this in 2009 with the introduction
of two marine protected areas. In the Philippines, the taking of mantas was banned in 1998, but this was
overturned in 1999 under pressure from local fishermen. Fish stocks were surveyed in 2002, and the ban was
reintroduced. The taking or killing of mantas in Mexican waters was prohibited in 2007. This ban may not be
strictly enforced, but laws are being more rigidly applied at Isla Holbox, an island off the Yucatán Peninsula,
where manta rays are used to attract tourists.

In 2009, Hawaii became the first of the United States to introduce a ban on the killing or capturing of manta
rays. Previously, no fishery for mantas existed in the state but migratory fish that pass the islands are now
protected. In 2010, Ecuador introduced a law prohibiting all fishing for manta and other rays, their retention as
bycatch and their sale.[17]

Relation with humans

The ancient Peruvian Moche people worshipped the sea and its animals. Their art often depicts manta rays.[56]
Historically, mantas were feared for their size and power. Sailors believed that they were dangerous to humans
and could sink boats by pulling on the anchors. This attitude changed around 1978, when divers around the
Gulf of California found them to be placid and that they could interact with the animals. Several divers
photographed themselves with mantas, including Jaws author Peter Benchley.[57]

Aquariums

Ceramic manta ray made by Moche
people, 200 AD, Larco Museum
Lima, Peru
ray joined that aquarium's collection in September 2009[59] and a third was added in 2010.[60]
Due to their size, mantas are
rarely kept in captivity and
few aquaria currently display
them. One notable individual
is "Nandi", a manta ray
which was accidentally
caught in shark nets off
Durban, South Africa, in
2007. Rehabilitated and Manta alfredi at Okinawa Churaumi
outgrowing her aquarium at Aquarium
uShaka Marine World, Nandi
was moved to the larger
Georgia Aquarium in August 2008, where she resides in its 23,848
m3 (6,300,000 US gal) "Ocean Voyager" exhibit.[58] A second manta

The Atlantis resort on Paradise Island, Bahamas, hosted a manta named "Zeus" that was used as a research
subject for 3 years until it was released in 2008.[61] The Okinawa Churaumi Aquarium also houses manta rays
in the "Kuroshio Sea" tank, one of the largest aquarium tanks in the world. The first manta ray birth in
captivity took place there in 2007. Although this pup did not survive, the aquarium has since had the birth of
three more manta rays in 2008, 2009, and 2010.[38]

Tourism

Sites at which manta rays congregate attract tourists, and manta

viewing generates substantial annual revenue for local
communities.[23]:19 Tourist sites exist in the Bahamas, the Cayman
Islands, Spain, the Fiji Islands, Thailand, Indonesia, Hawaii, Western
Australia[62] and the Maldives.[63] Mantas are popular because of
their enormous size and because they are easily habituated to humans.
Scuba divers may get a chance to watch mantas visiting cleaning
stations and night dives enable viewers to see mantas feeding on
Manta alfredi and scuba diver
plankton attracted by the lights.[64]

Ray tourism benefits locals

and visitors by raising awareness of natural resource management and
educating them about the animals.[62] It can also provide funds for
research and conservation.[63] Constant unregulated interactions with
tourists can negatively affect the fish by disrupting ecological
relationships and increasing disease transmission.[62] At Bora Bora,
an excessive number of swimmers, boaters, and jet skiers caused the
local manta ray population to abandon the area.[23]:19

In 2014, Indonesia brought in a fishing and export ban, as it has Play media
realized that manta ray tourism is more economically beneficial than M. alfredi during a dive at Hawaii
allowing the fish to be killed. A dead manta is worth $40 to $500,
while manta ray tourism can bring in $1 million during the life of a
single manta ray. Indonesia has 5.8 million km2 (2.2 million mi2 ) of ocean, and this is now the world's largest
sanctuary for manta rays.[65]

See also
 List of threatened rays
Manta Matcher
Manta Ray

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External links
Maldives Manta Rays: VIDEO (https://www.facebook.com/dcaquaventure/videos/17732410094
25587/)
Microdocs (http://www.stanford.edu/group/microdocs/): Manta ray (http://www.stanford.edu/grou
p/microdocs/mantaray.html)
Manta Ray videos and news stories from the BBC including footage of the possible new
species (https://web.archive.org/web/20091115103422/http://www.bbc.co.uk/nature/species/Ma
nta_ray)
Manta birostris, IUCN Red List of Threatened Species (https://web.archive.org/web/201904040
85817/http://oldredlist.iucnredlist.org/details/198921/0)
Manta ray (https://web.archive.org/web/*/www.arkive.org/) media from ARKive
 Diving with Mantas at the Azores (https://www.youtube.com/watch?v=h-TmonIb3MU)
Meet the Maldivian Mantas (https://web.archive.org/web/20130604113636/http://www.mantatru
st.org/photo-stories/meet-the-maldivian-mantas/) Manta Trust.
Manta Matcher – The Wildbook for Manta Rays (http://mantamatcher.org/)
Japans giant manta ray, Okinawa world’s first to exhibit (https://marinehobby.com/2018/12/09/ja
pans-giant-manta-ray-okinawa-worlds-first-to-exhibit/)

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