Diversity
Rays of the family Myliobatidae are well known for their extreme grace and great size.
With three subfamilies containing seven genera and about 42 species, the family includes
the eagle rays, manta or devil rays, and cownose rays (see Systematic/Taxonomic History).
These are free-swimming rays with broad, powerful pectoral fins that can measure over
6 m from tip to tip (see Physical Description). Many members of the family are able
to leap completely out of the water into the air. Rays, like their shark relatives
(both fall within the superorder
Euselachii
), have a reproductive strategy in which a great deal of energy is invested in relatively
few young over a lifetime, which may last several decades. Myliobatids generate, depending
on the subfamily, only one to six embryos each year, and these young are born live
after growing to considerable size inside the mother’s uterus (see Development, also
Reproduction). Despite the imposing size attained by many members of the family, these
rays are not dangerous to humans. Mantas have tiny teeth and strain planktonic organisms
(and sometimes small schooling fishes) from the water. Eagle and cownose rays have
pavement-like teeth suited for grinding mollusks (see Food Habits). Their tail spines,
when present, are used for defense. The worst damage caused by these rays is economic,
for they are capable of destroying entire beds of cultivated mollusks or oysters (see
Economic Importance to Humans). As of 1994 only one species was listed as vulnerable
to extinction, but due to their “slow” reproductive strategy rays may have difficulty
replenishing their numbers if human activity threatens them.
Geographic Range
Eagle, cownose, and manta or devil rays occur in tropical and warm temperate seas
worldwide. They are found in the Atlantic, Pacific, and Indian oceans, as well as
smaller seas. In the Atlantic some migrate as far north as the British Isles and Cape
Cod.
- Biogeographic Regions
- nearctic
- palearctic
- oriental
- ethiopian
- neotropical
- australian
- oceanic islands
- indian ocean
- atlantic ocean
- pacific ocean
- mediterranean sea
Habitat
Myliobatidae is a marine family, although some eagle and cownose venture into estuaries
and mangrove areas. Cosmopolitan in tropical and warm temperate seas, this family
can be found near reefs, in coastal lagoons, and, especially in the case of manta
rays, far out to sea. Many members of the family make summertime migrations into temperate
waters. Although cownose and eagle rays spend time feeding on mollusks and other invertebrates
on or near the substrate, they, along with the plankton-feeding manta rays, are free-swimming.
- Habitat Regions
- temperate
- tropical
- saltwater or marine
- Aquatic Biomes
- pelagic
- reef
- coastal
- brackish water
- Other Habitat Features
- estuarine
- intertidal or littoral
Systematic and Taxonomic History
Compagno (in Hamlett, 1999) notes that “there are at least as many classifications…of
cartilaginous fishes as authors who have published them,” and that “the higher classification
of chondrichthyans is currently in a state of flux.” Although there is less disagreement
regarding the classification of lower taxa (familial, generic and specific), there
are still a number of divergences in opinion at the familial level. Compagno (in Hamlett,
1999) maintains his earlier (1973) classification, in which Myliobatidae (eagle rays),
Rhinopteridae (cownose rays), and Mobulidae (manta or devil rays) each stand as a
separate family. Here we follow Nelson (1994), who acknowledges Compagno’s work but
places the three groups, as subfamilies, into the single family Myliobatidae. The
three subfamilies are Myliobatinae (eagle rays), with four genera and about 24 species;
Rhinopterinae (cownose rays) with one genus and five species; Mobulinae (manta rays
and devil rays) with two genera and 13 species.
Physical Description
In general, eagle, cownose, and manta or devil rays are medium to large, heavy-bodied
fishes with winglike pectoral fins forming a disc that is much wider than it is long.
The tail is much longer than the disc, has a filamentous end, and in many species
bears one or more serrated stinging spines near its base, close behind the pelvic
fins. In these rays the head is elevated and protrudes in front of the disc, but an
anterior portion of the pectoral fins continues forward to form a subrostral lobe
under the snout. In the subfamily Myliobatinae (eagle rays) this lobe is rounded.
The subfamily Rhinopterinae gets its common name, cownose ray, from the indentation
in the middle of the snout and forehead that forms a double fleshy lobe. In manta
or devil rays (subfamily Mobulinae) this lobe takes the form of two long, mobile,
cephalic fins, making manta rays the only living vertebrates with three pairs of functional
limbs. These can be rolled into a spiral while swimming or extended like a funnel
to guide planktonic organisms into the ray’s mouth. Mantas take in water through their
mouths and filter out organisms with a highly developed branchial sieve. Their mouths
are large and terminal or subterminal, and they have very small, cuspidate or flat-crowned
teeth. The other two subfamilies, cownose and eagle rays, have small, subterminal
mouths with one to seven rows of large, flat teeth that form pavement-like plates
for crushing mollusks. The middle rows of teeth are much wider than the outer rows.
Some common characteristics shared by all three subfamilies are the presence of a
moderately large dorsal fin over or just behind the pelvic fins, a very small or absent
caudal fin, eyes lateral on the head, and two spiracles (respiratory openings) on
top of the head just behind the eyes. Like other batoids, the lower eyelid is intact
while the upper eyelid is fused to the eyeball. There are five pairs of gill openings,
large in Mobulinae but small in Myliobatinae and Rhinopterinae. These rays are generally
countershaded, ventrally white or pale and dorsally black, olive, gray, or brown.
Some (certain species of eagle ray) have dorsal spots or bands. The disc may be naked
or may be covered with small denticles. All three subfamilies can be large fish, with
adults measuring between one and five m long. Manta rays (subfamily Mobulinae) are
the largest of all rays, with some attaining pectoral fin spans of over 6 m, and weighing
more than 1360 kg. Members of this subfamily are reported to be the morphologically
the most highly evolved of all living elasmobranch fishes.
The male sexual organs are called mixopterygia or claspers, and can be found at the
rear bases of the pelvic fins. There are two claspers, and each has a groove that
fills with sperm before the clasper is inserted into a female. Female eagle, cownose,
and manta rays have a uterus that is specially adapted to nurturing embryos outside
of the egg (see Development and Reproduction below).
- Other Physical Features
- ectothermic
- bilateral symmetry
- venomous
Development
Members of the family Myliobatidae, like other rays and their shark relatives, employ
a reproductive strategy that involves putting a great investment of energy into relatively
few young over a lifetime. One species,
Aetobatus narinari
, is known to reach sexual maturity at four to six years of age, but others may not
reach maturity until much older. Once sexually mature, rays have only one litter per
year, and in manta rays and some cownose rays a litter consists of only one embryo.
Since few young are produced, it is important that they survive, and to this end rays
are born at a large size, able to feed and fend for themselves much like an adult.
Rays develop from egg to juvenile inside the mother’s uterus, sometimes to almost
half their adult size. In this system, called aplacental uterine viviparity, developing
embryos receive most of their nutriment from a milky, organically rich substance secreted
by the mother’s uterine lining. An embryo absorbs this substance, called histotroph,
by ingestion, or through its skin or other specialized structures. Rays’ eggs are
small and insufficient to support the embryos until they are born, although the first
stage of development does happen inside tertiary egg envelopes that enclose each egg
along with egg jelly. The embryo eventually absorbs the yolk sac and stalk and the
histotroph provides it with nutrition, so much so that in
Rhinoptera bonasus
, for example, the embryo’s net weight increases to 3,000 times that of the egg. Development
from egg to term (birth) usually takes about two to four months. At birth the ray
is fully developed and the umbilicus completely absorbed.
Reproduction
Only a few species of elasmobranch (subclass including all sharks and rays) fishes
have been observed during courtship and mating. However, rays have a system that involves
internal fertilization, and elasmobranch fishes have relatively complex endocrine
(hormonal) systems. Based on knowledge of other vertebrates with similar systems,
it is likely that females signal to males through chemical or behavioral cues to indicate
when their hormonal state is appropriate for mating. Rays in general are reported
to engage in prolonged mating that may last up to four hours. According to one observation
of manta rays, mating occurred right at the surface of the water, with a graceful
undulating motion of their bodies, and the male alternately inserting his claspers
(paired male reproductive organs, also known as mixopterygia) into the female. The
pair did not copulate continuously, but swam about for short periods.
Rays bear young on a yearly cycle, although pregnancy usually lasts only several months,
generally spanning some period in the spring, summer, and fall. Only a few species,
like
Myliobatis californicus
with its nearly year-long gestation period, are known to differ from this pattern.
Within any given group of rays, individuals appear to go through mating, gestation,
and parturition (birth) at the same time as all the other females in the group. Eagle
rays bear up to six young at a time, but all manta rays and at least some cownose
rays bear only one pup in each cycle. A milky fluid (histotroph) secreted by the uterus
nourishes the embryos (see Development for a description of this system, called aplacental
uterine viviparity). In some groups the epithelium, or wall, of the uterus has evolved
to form trophonemata, elongated villi that extend into the uterine cavity to provide
greater surface area for respiratory exchange and histotroph excretion. This advanced
system of nourishing young inside the uterus can produce offspring that are relatively
large at birth (see Development). According to one investigator, a young ray is rolled
up like a cigar during birth, which, along with the lubricating histotroph, facilitates
the birth of such proportionally large young. The young ray then unrolls and swims
away. Likewise, sting-bearing young are able to pass out of the mother’s body without
stinging her because their stings are encased in a pliable sheath that sloughs off
after birth.
- Key Reproductive Features
- iteroparous
- seasonal breeding
- gonochoric/gonochoristic/dioecious (sexes separate)
- sexual
- fertilization
- viviparous
No reported evidence of parental care in Myliobatidae was found. After such extended nurturing inside their mothers’ bodies, young rays come into the sea quite able to feed and fend for themselves (see Development and Reproduction).
- Parental Investment
- no parental involvement
Lifespan/Longevity
There was little specific information information found regarding lifespans in Myliobatidae,
but in general rays, like their relatives the sharks, grow and mature slowly and are
long-lived. It is known that
spotted eagle rays
reach sexual maturity after four to six years, and
bat rays
live about 23 years. Some researchers estimate that the largest sharks and rays may
not reach maturity until 20 to 30 years of age, and that they may live to maximum
ages of 70 to 100 years or more.
Behavior
Eagle, manta, and cownose rays are more active than their bottom-dwelling stingray
relatives. They swim freely in the water column, near reefs or over a continental
shelf, often near the surface. Eagle rays occur alone, in pairs, or in schools of
a hundred or more. Cownose rays have been observed in groups of 4000 to 6000, apparently
migrating. Manta or devil rays, although frequently observed singly, also occur in
small shoals of up to five individuals. Members of this family are famous for their
ability to leap high into the air, sometimes turning a somersault. Some hypothesize
that this activity serves to rid the rays of parasites, while others view the behavior
as playful. Like other rays, they use their broad pectoral fins rather than the tail
for forward propulsion. This results in their graceful “flying” motion through the
water. Manta rays have been observed gliding near the surface with the upturned tips
of their pectoral fins sticking up out of the water.
Remoras
often accompany manta rays, inside their mouths or clinging to their pectoral fins.
Eagle rays seem to travel around in search of mollusks, which they dig up, using their
snouts like a spade. Cownose rays often feed in schools, beating their powerful pectoral
fins to expose buried shellfish.
- Key Behaviors
- natatorial
- diurnal
- motile
- migratory
- social
Communication and Perception
Rays perceive and interact with their environment using sensory channels common to
many vertebrates: sight, hearing, smell, taste and touch. Rays also belong to a group
of fishes, the elasmobranchs, whose electrical sensitivity seems to exceed that of
all other animals. Elasmobranch fishes are equipped with ampullae of Lorenzini, electroreceptor
organs that contain receptor cells and canals leading to pores in the animal’s skin.
Sharks and rays can detect the electrical patterns created by nerve conduction, muscular
contraction, and even the ionic difference between a body (i.e. of prey) and water.
In lab experiments, stingrays changed their feeding location according to artificially
induced changes in the electrical field around them. Other experiments have demonstrated
that cartilaginous fishes use electrosensory information not only to locate prey,
but also for orientation and navigation based on the electrical fields created by
the interaction between water currents and the earth’s magnetic field. Although some
rays can produce an electric shock to defend themselves or stun prey, members of the
family Myliobatidae cannot. Some of them are able, however, to inflict a venomous
sting with their tail spine in defense. Rays sometimes let out a sound like a bark
when captured.
Food Habits
Members of the subfamily Mobulinae, the manta and devil rays, strain their food from
the water. The large cephalic fins on either side of the mouth (see Physical Description)
can be extended to form a funnel that scoops planktonic organisms, small fishes and
crustaceans into the ray’s mouth. Manta rays have a highly developed filter system
on their gill arches that traps small animals taken in with water during respiration.
The other two subfamilies, Myliobatinae (eagle rays) and Rhinopterinae (cownose rays),
have interlocking plates of flat, pavement-like teeth suited for grinding mollusks
and other hard-shelled prey. Some, however, have also been observed eating fish, octopus,
and worms.
Spotted eagle rays
are able to crush clams and oysters and spit out the shells, accomplishing this so
deftly that whole soft oysters, without the shell, have been found in the rays’ stomachs.
Cownose and eagle rays use their powerful pectoral fins to fan the substrate, creating
a suction that digs out buried clams, and then use the lower parts of their snouts
to pry up the mollusks. All members of the family Myliobatidae appear to move around
in search of concentrations of prey.
- Foraging Behavior
- filter-feeding
Predation
Although rays can grow very large, they are still preyed upon by other large fishes,
especially sharks. The great hammerhead
Sphyrna mokarran
, in particular, appears to specialize in eating stingrays. It uses its hammer head
to knock a ray to the bottom, and then pins the ray, once again with its head, pivoting
around to bite the ray’s disc until the ray succumbs and can be eaten.
Ecosystem Roles
Eagle, manta, and cownose rays are nearly cosmopolitan in tropical and warm temperate
seas, and therefore are a consistent predator on populations of mollusks, crustaceans,
planktonic organisms, and small fishes. They, especially smaller specimens, provide
food for sharks and other large fishes.
Economic Importance for Humans: Positive
While little information was found regarding human use of the family Myliobatidae
in particular, cartilaginous fishes in general are important to humans in a number
of ways. Australian Aborigines have eaten rays for centuries. They determine whether
a seasonal catch is ready to eat by checking a ray’s liver; if it is oily and pinkish
white, the ray is suitable for eating. Manta rays, however, along with rays that have
two spines, are considered inedible. Aborigines use ray spines for spear tips, although
this likely impacts stingrays of other families more than members of Myliobatidae.
Rays are considered food fish in Australia, Europe, and parts of Asia, and in some
places are among the most highly priced fishes. Like shark fins, fins of some rays
are harvested in Asia for soup and as an aphrodisiac. Cartilaginous fishes are used
for medical purposes as well. Chondroiten, used as skin replacement for burn victims,
is derived from the fishes’ cartilage. Other extracts from cartilage help suppress
tumors and may assist cancer treatment. Finally, some large rays are a popular part
of public aquarium exhibits.
- Positive Impacts
- food
- source of medicine or drug
Economic Importance for Humans: Negative
Eagle rays and cownose rays are considered a nuisance wherever humans cultivate oyster
or clam beds. They are capable of clearing an entire area of mollusks, and destroy
the beds. If these rays perceive the need to defend themselves, they can deliver a
stab with their tail spine that is reported to be excruciating. Members of Myliobatidae,
however, cannot whip their tails about as effectively as other stingrays because their
sting is located near the base of the tail. The largest members of the family, the
manta rays, are plankton feeders and are harmless to humans and their commercial endeavors.
- Negative Impacts
-
injures humans
- bites or stings
- venomous
- crop pest
Conservation Status
As of 1994 only one species,
giant devil ray
, was listed as vulnerable to extinction. However, sharks and rays in general are
vulnerable to overfishing. They grow and mature slowly, and the size of the adult
population closely determines the number of young produced, due to their “slow” reproductive
strategy of investing a great deal of energy in relatively few young over a lifetime.
Other Comments
Myliobatidae is a family of cartilaginous fishes (class
Chondrichthyes
), the oldest surviving group of jawed vertebrates. This group was the first to bear
live young, nourish developing embryos by means of a placenta, and to regulate reproduction
and embryonic growth hormonally. Batoids (skates and rays) split off from the sharks
in the early Jurassic period. Fossil records of Myliobatidae date back to the upper
Cretaceous period.
Additional Links
Contributors
Monica Weinheimer (author), Animal Diversity Web.
R. Jamil Jonna (author), Animal Diversity Web.
- Nearctic
-
living in the Nearctic biogeographic province, the northern part of the New World. This includes Greenland, the Canadian Arctic islands, and all of the North American as far south as the highlands of central Mexico.
- native range
-
the area in which the animal is naturally found, the region in which it is endemic.
- Palearctic
-
living in the northern part of the Old World. In otherwords, Europe and Asia and northern Africa.
- native range
-
the area in which the animal is naturally found, the region in which it is endemic.
- oriental
-
found in the oriental region of the world. In other words, India and southeast Asia.
- native range
-
the area in which the animal is naturally found, the region in which it is endemic.
- Ethiopian
-
living in sub-Saharan Africa (south of 30 degrees north) and Madagascar.
- native range
-
the area in which the animal is naturally found, the region in which it is endemic.
- Neotropical
-
living in the southern part of the New World. In other words, Central and South America.
- native range
-
the area in which the animal is naturally found, the region in which it is endemic.
- Australian
-
Living in Australia, New Zealand, Tasmania, New Guinea and associated islands.
- native range
-
the area in which the animal is naturally found, the region in which it is endemic.
- oceanic islands
-
islands that are not part of continental shelf areas, they are not, and have never been, connected to a continental land mass, most typically these are volcanic islands.
- native range
-
the area in which the animal is naturally found, the region in which it is endemic.
- native range
-
the area in which the animal is naturally found, the region in which it is endemic.
- Atlantic Ocean
-
the body of water between Africa, Europe, the southern ocean (above 60 degrees south latitude), and the western hemisphere. It is the second largest ocean in the world after the Pacific Ocean.
- native range
-
the area in which the animal is naturally found, the region in which it is endemic.
- Pacific Ocean
-
body of water between the southern ocean (above 60 degrees south latitude), Australia, Asia, and the western hemisphere. This is the world's largest ocean, covering about 28% of the world's surface.
- native range
-
the area in which the animal is naturally found, the region in which it is endemic.
- native range
-
the area in which the animal is naturally found, the region in which it is endemic.
- temperate
-
that region of the Earth between 23.5 degrees North and 60 degrees North (between the Tropic of Cancer and the Arctic Circle) and between 23.5 degrees South and 60 degrees South (between the Tropic of Capricorn and the Antarctic Circle).
- tropical
-
the region of the earth that surrounds the equator, from 23.5 degrees north to 23.5 degrees south.
- saltwater or marine
-
mainly lives in oceans, seas, or other bodies of salt water.
- pelagic
-
An aquatic biome consisting of the open ocean, far from land, does not include sea bottom (benthic zone).
- reef
-
structure produced by the calcium carbonate skeletons of coral polyps (Class Anthozoa). Coral reefs are found in warm, shallow oceans with low nutrient availability. They form the basis for rich communities of other invertebrates, plants, fish, and protists. The polyps live only on the reef surface. Because they depend on symbiotic photosynthetic algae, zooxanthellae, they cannot live where light does not penetrate.
- coastal
-
the nearshore aquatic habitats near a coast, or shoreline.
- brackish water
-
areas with salty water, usually in coastal marshes and estuaries.
- estuarine
-
an area where a freshwater river meets the ocean and tidal influences result in fluctuations in salinity.
- intertidal or littoral
-
the area of shoreline influenced mainly by the tides, between the highest and lowest reaches of the tide. An aquatic habitat.
- ectothermic
-
animals which must use heat acquired from the environment and behavioral adaptations to regulate body temperature
- bilateral symmetry
-
having body symmetry such that the animal can be divided in one plane into two mirror-image halves. Animals with bilateral symmetry have dorsal and ventral sides, as well as anterior and posterior ends. Synapomorphy of the Bilateria.
- venomous
-
an animal which has an organ capable of injecting a poisonous substance into a wound (for example, scorpions, jellyfish, and rattlesnakes).
- iteroparous
-
offspring are produced in more than one group (litters, clutches, etc.) and across multiple seasons (or other periods hospitable to reproduction). Iteroparous animals must, by definition, survive over multiple seasons (or periodic condition changes).
- seasonal breeding
-
breeding is confined to a particular season
- sexual
-
reproduction that includes combining the genetic contribution of two individuals, a male and a female
- fertilization
-
union of egg and spermatozoan
- internal fertilization
-
fertilization takes place within the female's body
- viviparous
-
reproduction in which fertilization and development take place within the female body and the developing embryo derives nourishment from the female.
- natatorial
-
specialized for swimming
- diurnal
-
- active during the day, 2. lasting for one day.
- motile
-
having the capacity to move from one place to another.
- migratory
-
makes seasonal movements between breeding and wintering grounds
- social
-
associates with others of its species; forms social groups.
- visual
-
uses sight to communicate
- acoustic
-
uses sound to communicate
- visual
-
uses sight to communicate
- tactile
-
uses touch to communicate
- acoustic
-
uses sound to communicate
- chemical
-
uses smells or other chemicals to communicate
- electric
-
uses electric signals to communicate
- magnetic
-
(as perception channel keyword). This animal has a special ability to detect the Earth's magnetic fields.
- filter-feeding
-
a method of feeding where small food particles are filtered from the surrounding water by various mechanisms. Used mainly by aquatic invertebrates, especially plankton, but also by baleen whales.
- food
-
A substance that provides both nutrients and energy to a living thing.
- drug
-
a substance used for the diagnosis, cure, mitigation, treatment, or prevention of disease
- venomous
-
an animal which has an organ capable of injecting a poisonous substance into a wound (for example, scorpions, jellyfish, and rattlesnakes).
- carnivore
-
an animal that mainly eats meat
- piscivore
-
an animal that mainly eats fish
References
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Allen, T. 1996. Shadows in the Sea: The Sharks, Skates, and Rays . New York, NY: Lyons and Buford.
Berg, L. 1958. System Der Rezenten und Fossilen Fischartigen und Fische . Berlin: VEB Deutscher Verlag der Wissenschaften.
Bleckmann, H., M. Hofmann. 1999. Special Senses. Pp. 300-328 in Sharks, Skates, and Rays . Baltimore, MD: The Johns Hopkins University Press.
Böhlke, J., C. Chaplin. 1968. Fishes of the Bahamas and Adjacent Tropical Waters . Wynnewood, PA: Published for the Academy of Natural Sciences of Philadelphia by Livingston.
Compagno, L. 1999. Systematics and Body Form. Pp. 1-42 in Sharks, Skates, and Rays . Baltimore, MD: The Johns Hopkins University Press.
Hamlett, W., T. Koob. 1999. Female Reproductive System. Pp. 398-443 in Sharks, Skates, and Rays . Baltimore, MD: The Johns Hopkins University Press.
Hamlett, W. 1999. Male Reproductive System. Pp. 444-470 in Sharks, Skates, and Rays . Baltimore, MD: The Johns Hopkins University Press.
Helfman, G., B. Collete, D. Facey. 1997. The Diversity of Fishes . Malden, MA: Blackwell.
Last, P., J. Stevens. 1994. Sharks and Rays of Australia . Australia: CSIRO.
Moyle, P., J. Cech. 2000. Fishes: An introduction to ichthyology – fourth edition . Upper Saddle River, NJ: Prentice-Hall.
Nelson, J. 1994. Fishes of the World – third edition . New York, NY: John Wiley and Sons.
The World Conservation Union, 2002. "IUCN 2002" (On-line). 2002 IUCN Red List of Threatened Species. Accessed October 15, 2003 at http://www.redlist.org/ .
Wheeler, A. 1985. The World Encyclopedia of Fishes . London: Macdonald.
Wourms, J., L. Demski. 1993. The reproduction and development of sharks, skates, rays, and ratfishes: introduction, history, overview, and future prospects. Pp. 7-19 in The Reproduction and Development of Sharks, Skates, Rays, and Ratfishes . Dordrecht, The Netherlands: Kluwer Academic Publishers.