Diversity
The family Gymnuridae contains two genera and about 12 species. Their pectoral fins
form a disc that is much broader than it is long, forming “wings” that give the family
its common name, butterfly rays. These rays are marine and occasionally estuarine,
found worldwide in tropical and subtropical seas. They live on the bottom in open
sandy areas, where they feed on a variety of invertebrates, fishes, and other small
organisms. Some bear a venomous spine on their short tails, but butterfly rays are
considered far less dangerous than the long-tailed stingrays. Like other rays, butterfly
rays are viviparous, giving birth to live young after nourishing them inside the uterus
(see Development and Reproduction).
Geographic Range
Members of the family Gymnuridae can be found worldwide in tropical and warm temperate
seas. Their range in the Atlantic includes the Black Sea, and they are widely distributed
in the Pacific and Indian oceans.
- Biogeographic Regions
- nearctic
- palearctic
- oriental
- ethiopian
- neotropical
- australian
- oceanic islands
- indian ocean
- atlantic ocean
- pacific ocean
- mediterranean sea
Habitat
Gymnuridae is a marine family, but are occasionally found in estuaries and river mouths
as well. Butterfly rays are benthic (bottom-dwellers), most commonly found on the
upper continental shelf. They prefer stretches of open sand and frequent shallow beaches
and bays.
- Habitat Regions
- temperate
- tropical
- saltwater or marine
- Aquatic Biomes
- benthic
- coastal
- brackish water
- Other Habitat Features
- estuarine
- intertidal or littoral
Physical Description
As their name indicates, butterfly rays have extremely wide pectoral fins that resemble
butterfly wings. These fins create a disc that is more than 1.5 times as broad as
it is long and extends forward to include the head, ending in a blunt, angular, short
snout. The snout is rounder and blunter on females than on males, at least in the
case of
Gymnura micrura
. The disc is brown, grayish, purplish or greenish in color, sometimes with pale and
dark spots, and it may be naked or covered with small denticles. The head is even
with the disc rather than elevated, giving the fish a very flat appearance from the
side view. The eyes are located on the sides of the top of the head, with the spiracles
(respiratory openings) close behind the eyes. At least one species,
Gymnura australis
, has a long tentacle near the back edge of the spiracles. Like all rays, butterfly
rays have ventral gill openings. These form five small pairs and the internal gill
arches do not have filter plates. Their mouths are relatively large and located under
the end of the snout. Since their mouths are directed downward and often placed against
the sand, bottom-living rays use their spiracles rather than their mouths for water
intake, and, if the gills are covered with sand, the spiracles are also used for expelling
water. Butterfly rays have small, cuspidate teeth that do not form the crushing plates
found in many other rays. Their dorsal fins, if present, are small. They lack anal
fins, and rather than a caudal fin they may have low dorsal and ventral ridges on
the tail. The slender tail is much shorter than the disc. Some butterfly rays have
a stinging spine behind the pelvic fins. These fish range in size from less than 0.5
m long to over 2 m long.
- Other Physical Features
- ectothermic
- bilateral symmetry
- venomous
- Sexual Dimorphism
- sexes alike
- sexes shaped differently
Development
Members of the family Gymnuridae, 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. 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. Researchers have found that in some stingrays, the stomach and spiral
intestine are among the first organs to develop and function, so that the embryo can
digest the uterine “milk.” 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. Embryos are
so well nourished in the uterus that in
Gymnura micrura
, for example, the young ray’s net weight increases by 4900% from egg to birth, and
are between 15-22 cm when they are born. Development in the uterus usually takes about
two to four months. One species of butterfly ray,
Gymnura natalensis
, is born with an almost triangular shape, but develops much wider pectoral fins as
it grows, and in adults the disc is more than twice as broad as it is long.
Reproduction
Only a few species of elasmobranch (subclass including all sharks and rays) fishes
have been observed during courtship and mating. However, butterfly rays have a system
that involves internal fertilization, so it can logically be inferred that mating
communication between male and female must happen to an extent that allows the male
to insert at least one of his two claspers (male reproductive organs that are modifications
of the pelvic fins) into the female’s cloaca to deposit sperm. 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 bear young on a yearly cycle, although pregnancy usually lasts only several months,
generally spanning some period in the spring, summer, and fall. 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. Butterfly rays usually bear
between two and six live young at a time, after nourishing the embryos with milky
fluid (histotroph) secreted by the uterus (see Development for a description of this
system, called aplacental uterine viviparity). In Gymnuridae 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 passage 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 Gymnuridae 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
Little specific information regarding lifespans in Gymnuridae was found, but in general
rays, like their relatives the sharks, grow and mature slowly and are long-lived.
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. The family Gymnuridae does not include the largest rays and may not reach
such extremes of longevity.
Behavior
At least one species of butterfly ray (
Gymnura micrura
) is migratory, moving into temperate waters during the warm season. Butterfly rays
forage for food over the sandy bottoms on which they live. Some butterfly rays are
equipped with a venomous tail spine that they use in defense. Due to their short tails
and smaller spines, however, their stings are generally less fierce than those of
their stingray relatives.
- Key Behaviors
- natatorial
- motile
- migratory
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 Gymnuridae cannot. Some are able, however, to inflict a sting with their tail
spine in defense.
Food Habits
Butterfly rays feed upon crabs, shrimps, various invertebrates, fishes, small crustaceans,
and mollusks. They generally seek prey near the bottom of warm waters over sandy stretches.
- Primary Diet
-
carnivore
- piscivore
- eats non-insect arthropods
- molluscivore
Predation
Although rays can grow very large, they are still preyed upon by other large fishes,
especially sharks. Stingray spines have been found embedded in the mouths of many
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. In addition
to their defensive venomous sting, many rays have drab coloring that matches the sand
or mud bottom. Several species of butterfly ray can quickly change color to blend
in with their background.
- Anti-predator Adaptations
- cryptic
Ecosystem Roles
Stingrays are nearly cosmopolitan in tropical and warm temperate seas, and therefore
are a consistent predator on populations of mollusks, crustaceans, worms, and fishes.
They, in turn, provide food for sharks and other large fishes.
Remoras
sometimes accompany adult rays.
Economic Importance for Humans: Positive
Although butterfly rays do not have any particular commercial importance, 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. Rays that have two spines, however, are considered inedible. Australian
Aborigines, Malayans, tribes in South and Central America, and West Africa, and peoples
of the Indo-Pacific have used ray spines for spear tips, daggers, or whips. Rays are
considered food fish in Australia, Europe, and parts of Asia, and in some places are
among the most highly priced fishes. In Australia, cuts from one species of butterfly
ray are sometimes sold as skate flesh. 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. Some large rays are a popular part of public aquarium
exhibits.
- Positive Impacts
- food
Economic Importance for Humans: Negative
Butterfly rays have little negative impact on humans. They are not known to destroy
shellfish beds, and their spines, when present, are much less dangerous than those
of their long-tailed relatives (
Dasyatidae
).
- Negative Impacts
-
injures humans
- bites or stings
- venomous
Conservation Status
Currently, there is no conservation threat to any member of this family.
Additional Links
- 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.
- 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.
- benthic
-
Referring to an animal that lives on or near the bottom of a body of water. Also an aquatic biome consisting of the ocean bottom below the pelagic and coastal zones. Bottom habitats in the very deepest oceans (below 9000 m) are sometimes referred to as the abyssal zone. see also oceanic vent.
- 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
- motile
-
having the capacity to move from one place to another.
- migratory
-
makes seasonal movements between breeding and wintering grounds
- visual
-
uses sight to communicate
- tactile
-
uses touch to communicate
- chemical
-
uses smells or other chemicals 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.
- cryptic
-
having markings, coloration, shapes, or other features that cause an animal to be camouflaged in its natural environment; being difficult to see or otherwise detect.
- food
-
A substance that provides both nutrients and energy to a living thing.
- 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
- molluscivore
-
eats mollusks, members of Phylum Mollusca
References
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Allen, T. 1996. Shadows in the Sea: The Sharks, Skates, and Rays . New York, NY: Lyons and Buford.
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.
Liem, K., A. Summers. 1999. Muscular System: Gross Anatomy and Functional Morphology of Muscles. Pp. 93-114 in Sharks, Skates, and Rays . Baltimore, MD: The Johns Hopkins University Press.
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, 2003. "IUCN 2002" (On-line). 2002 IUCN Red List of Threatened Species. Accessed November 21, 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. 7-21 in The Reproduction and Development of Sharks, Skates, Rays, and Ratfishes . Dordrecht, The Netherlands: Kluwer Academic Publishers.
Wourms, J. 1993. Maximization of evolutionary trends for placental viviparity in the spadenose shark, Scoliodon laticaudus. Pp. 269-293 in The Reproduction and Development of Sharks, Skates, Rays, and Ratfishes . Dordrecht, The Netherlands: Kluwer Academic Publishers.