Diversity
The Belontiidae family, commonly known as gouramies, is composed of approximately
12 genera and 46 species, and is the largest and most diverse family in the
Anabantoidei
suborder. Gouramies are restricted to fresh water and are capable of inhabiting stagnant,
low-oxygen areas due to the existence of an air-breathing organ called the labyrinthine
(in Physical Description below). In addition, the Belontiidae family contains some
of the most popular fish in the aquaria trade. One species,
Betta splendens
(
Siamese fighting fish
), has been in cultivation for over 100 years and captive individuals scarcely resemble
the wild variety (see Economic Importance for Humans). Due to their pugnacious behavior,
many gouramies have been used in behavioral studies as well (see Behavior).
Geographic Range
All members of the Belontiidae family inhabit freshwater and are indigenous to Africa
and Southern Asia. They can be found in Pakistan, India, China, Korea to the Malay
archipelago, Indonesia and Borneo. However, due to their popularity in the aquarium
trade and ease of transport, belontiids are frequently shipped far beyond their natural
range. Several species that now live in the Philippines and United States originally
escaped from the aquarium trade (Lever 1996 from Berra 2001 pg. 480, 487).
- Biogeographic Regions
- nearctic
- palearctic
- oriental
- ethiopian
Habitat
Gouramies are uniquely adapted to the stagnant waters of tropical areas. They inhabit
swift streams, the backwaters of large rivers, brackish lagoons, and potholes. They
are most common in stagnant areas with dense aquatic or overhanging vegetation. With
the ability to breathe air directly from the atmosphere, gouramies are able survive
in marginal to anoxic waters.
- Habitat Regions
- tropical
- freshwater
- Aquatic Biomes
- lakes and ponds
- rivers and streams
- temporary pools
- Other Habitat Features
- urban
- suburban
- agricultural
- riparian
Systematic and Taxonomic History
Liem (1963) established three subfamilies, representing the major evolutionary lines
of the Belontiidae family:
Belontiinae
(combtail gouramies), with one genus and two species,
Magropodinae
(Siamese fighting fishes, paradisefishes), with seven genera and approximately 32
species, and
Trighogastrinae
(gouramies), with four genera and about 13 species.
Gouramies are characterized by a distinct physical adaptation to stagnant waters,
which is the labyrinthine organ. In this organ, located above the gill chamber, intricate,
vascularized tissue stores air and allows gouramies to gulp air directly from the
atmosphere. Additionally, gouramies have a long body cavity, allowing the swimbladder
to extend into the tail.
Physical Description
Most gouramies are small (10 cm or less) and have elongate and cylindrical bodies.
Others are deep-bodied and compressed. Gouramies have rounded tails and the anal fin
base is much longer than the dorsal fin base in most species. In some gouramies, the
first ray on the pelvic fin is elongated and is used as a tactile (touch-sensing)
organ. After extensive breeding in captivity, the color and fin form have changed
considerably in some gouramies.
Sexual dimorphism
has been reported for some members of this family.
One of two distinct physical features is the labyrinthine organ located above the
gill chamber, which allows gouramies to breathe atmospheric air by capturing air bubbles
from the surface water and holding them in the labyrinthine organ until they are expelled
through the gill covers. “This unique organ is formed of highly vascularized, convoluted
tissue, supported by an enlarged dorsal element of the gill arches” (Johnson, G.D.
and A.C. Gill 1998). The bubbles taken in by gouramies are also important in hearing.
Air bubbles are stored adjacent to membranous windows on either side of the cranium,
which contains the inner ear. Slight vibrations picked up by the air bubble are easily
transmitted to the inner ear through this membranous window. The second feature is
a long body cavity that allows the swimbladder to extend into the caudal region. (Click
here to see a
fish diagram
).
- Other Physical Features
- ectothermic
- bilateral symmetry
- Sexual Dimorphism
- sexes colored or patterned differently
- sexes shaped differently
Development
After spawning, female gouramies release their eggs into the water. The eggs are gathered
by the male and left in a nest near the surface until fry hatch.
Reproduction
Female gouramies use pheromones to attract males during spawning. The female deposits
eggs into the water and they are collected by the male to deposit into the nest. The
courting behavior of some gouramies is quite elaborate and spawning territory is maintained
aggressively by the male.
Gouramies use air bubbles in constructing their nests. They create a nest of froth
by expelling mucous-covered bubbles at the surface, often beneath leaves. After gathering
the eggs, the male deposits them into the froth nest. The nest has a dual purpose:
to keep the developing young close together for protection and to keep them near the
water’s surface where it is well oxygenated.
- Key Reproductive Features
- iteroparous
- gonochoric/gonochoristic/dioecious (sexes separate)
- fertilization
- oviparous
Male gouramies guard the froth nest and maintain it by producing new bubbles as older
ones break down. If juveniles try to leave the nest early, the male transports them
back by carrying them in his mouth and spitting them back. In some genera (
Macropodus
and
Betta
) mouthbrooding has been reported as well.
- Parental Investment
- male parental care
Lifespan/Longevity
No specific information was found concerning lifespan or longevity for this family.
Behavior
Members of the
Betta
(
fighting gouramies
) genus have been used extensively in behavioral studies. Male fighting gouramies
are extremely belligerent toward each other and they are often bred to fight, as with
the fighting cocks.
Talking gouramies
use the air-breathing organ (labyrinthine) to produce a croaking sound, which is
used in establishing territories and dominance hierarchies. Additionally, some gouramies
use an elongated pelvic fin ray in exploratory behavior. The ray is directed forward
to gather information about the fish’s surroundings or sense another individual as
it approaches. Another interesting behavior of gouramies is the socially controlled
synchronous air breathing. Groups of individuals in both
Trichogaster
and
Colisa
come to the surface and gulp air simultaneously in a temporary school to lessen the
chance of predation while surfacing.
- Key Behaviors
- natatorial
- motile
- aestivation
- territorial
- social
- dominance hierarchies
Communication and Perception
Gouramies use several forms of communication:
Talking gouramies
use croaking to communicate, a modified pelvic fin ray is used as a tactile organ,
and chemical signals are used during mating. The former two are described in more
detail in the Behavior section and the latter is discussed in Reproduction.
- Other Communication Modes
- choruses
- pheromones
Food Habits
Members of the genera
Belontia
,
Trichopsis
, and
Macropodus
are omnivores, while
Trichogaster
is primarily herbivorous, and
Betta
and
Colisa
are carnivorous, feeding on shrimp, fish and insects in or out of water.
Colisa
uses the same technique as
archerfishes
to prey on insects just above the water. Bullets of water are created by compressing
the gill covers and forming a groove using the tongue and palate. The fish “shoots”
insects out of overhanging vegetation with these bullets and eats them when they hit
the water.
- Primary Diet
-
carnivore
- piscivore
- eats eggs
- insectivore
- eats non-insect arthropods
- herbivore
- omnivore
Predation
One impact of predation on gouramies is the frequency and location of atmospheric
air breathing. In the presence of predatory birds or fish, some gouramies hide in
mats of aquatic vegetation and surface less frequently. Other gouramies form temporary
schools when they come up in order to deter predators. This “synchronous air breathing”
is discussed above in Behavior.
Ecosystem Roles
Because gouramies are uniquely adapted to harsh conditions, such as hypoxia (water
without oxygen) and desiccation, they are an important predator in the stagnant and
intermittent waters of Asia. In addition, air breathing serves to protect gouramies
from excessive water pollution so they play an important role in the biological control
of insects in urban areas. This is discussed further in Economic Importance for Humans.
Economic Importance for Humans: Positive
Many gouramies are popular aquarium fishes because of their bright coloration, small
size, and interesting reproductive behaviors. Members of the genus
Betta
, fighting fishes, have been used extensively in genetic and behavioral studies. Fighting
fishes are also bred for entertainment and gambling in some countries. Finally, members
of the genus
Trichogaster
have been cultured in ponds as food fish.
Experiments with
Macropodus cupanus
show that breathing air at the surface instead of consuming oxygen entirely from
water lessens the absorption of toxic pollutants, which makes gouramies more effective
in the biological control of insects. For example, mosquito larvae form the main diet
of fish in the
Betta
genus. Each adult consumes approximately 10,000 to 15,000 mosquito larvae each year,
significantly reducing the mosquito populations and mosquito-related illnesses.
Economic Importance for Humans: Negative
No specific information was found concerning any negative impacts to humans.
Conservation Status
There are 11 species of concern within the genus
Betta
. Of these, three are listed as critically endangered, one is listed as endangered,
and seven are listed as vulnerable. Additionally,
Parosphromenus harveyi
is listed as endangered, and
Belontia signata
and
Malpulutta kretseri
are listed as low risk depending on conservation efforts.
Additional Links
Contributors
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.
- introduced
-
referring to animal species that have been transported to and established populations in regions outside of their natural range, usually through human action.
- 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.
- introduced
-
referring to animal species that have been transported to and established populations in regions outside of their natural range, usually through human action.
- 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.
- tropical
-
the region of the earth that surrounds the equator, from 23.5 degrees north to 23.5 degrees south.
- freshwater
-
mainly lives in water that is not salty.
- urban
-
living in cities and large towns, landscapes dominated by human structures and activity.
- suburban
-
living in residential areas on the outskirts of large cities or towns.
- agricultural
-
living in landscapes dominated by human agriculture.
- riparian
-
Referring to something living or located adjacent to a waterbody (usually, but not always, a river or stream).
- 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.
- 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).
- fertilization
-
union of egg and spermatozoan
- external fertilization
-
fertilization takes place outside the female's body
- oviparous
-
reproduction in which eggs are released by the female; development of offspring occurs outside the mother's body.
- male parental care
-
parental care is carried out by males
- natatorial
-
specialized for swimming
- motile
-
having the capacity to move from one place to another.
- territorial
-
defends an area within the home range, occupied by a single animals or group of animals of the same species and held through overt defense, display, or advertisement
- social
-
associates with others of its species; forms social groups.
- dominance hierarchies
-
ranking system or pecking order among members of a long-term social group, where dominance status affects access to resources or mates
- visual
-
uses sight to communicate
- tactile
-
uses touch to communicate
- acoustic
-
uses sound to communicate
- chemical
-
uses smells or other chemicals to communicate
- choruses
-
to jointly display, usually with sounds, at the same time as two or more other individuals of the same or different species
- pheromones
-
chemicals released into air or water that are detected by and responded to by other animals of the same species
- visual
-
uses sight to communicate
- tactile
-
uses touch to communicate
- acoustic
-
uses sound to communicate
- chemical
-
uses smells or other chemicals to communicate
- pet trade
-
the business of buying and selling animals for people to keep in their homes as pets.
- food
-
A substance that provides both nutrients and energy to a living thing.
- carnivore
-
an animal that mainly eats meat
- piscivore
-
an animal that mainly eats fish
- insectivore
-
An animal that eats mainly insects or spiders.
- herbivore
-
An animal that eats mainly plants or parts of plants.
- omnivore
-
an animal that mainly eats all kinds of things, including plants and animals
References
Berra, T. 2001. Freshwater Fish Distribution . San Diego, CA: Academic Press.
Froese, R., D. Pauly, D. Woodland. 2003. "Fish Base" (On-line). FishBase World Wide Web electronic publication. Accessed May 05, 2003 at http://www.fishbase.org/ .
Graham, J. 1997. Air-breathing fishes: evolution, diversity, and adaptation . San Diego, CA: Academic Press.
Helfman, G., B. Collete, D. Facey. 1997. The Diversity of Fishes . Malden, MA: Blackwell.
Johnson, G., A. Gill. 2002. Perches and Their Allies. Pp. 193 in Encyclopedia of Fishes – second edition . San Diego, CA: Academic Press.
McKinnon, J., N. Liley. 1987. Asymmetric species specificity in responses to female sexual pheromone by males of two species of Trichogaster (Pisces: Belontiidae). Can. Journal Zool. Dep. Zool., Univ. British Columbia, Vancouver, B.C. , 65 no. 5: 1129-1134.
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 July 08, 2003 at http://www.iucnredlist.org/ .
Wheeler, A. 1985. The World Encyclopedia of Fishes . London: Macdonald.