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The Mozambique tilapia (''Oreochromis mossambicus'') is an
oreochromine Oreochromini is a tribe of cichlids in the Pseudocrenilabrinae subfamily that is native to Africa and Western Asia, but a few species have been widely introduced to other parts of the world. It was formerly considered to be part of the tribe Ti ...
cichlid Cichlids () are a large, diverse, and widespread family of percomorph fish in the family Cichlidae, order Cichliformes. At least 1,760 species have been scientifically described, making it one of the largest vertebrate families, with on ...
fish native to southeastern Africa. Dull colored, the Mozambique tilapia often lives up to a decade in its native habitats. It is a popular fish for
aquaculture Aquaculture (less commonly spelled aquiculture), also known as aquafarming, is the controlled cultivation ("farming") of aquatic organisms such as fish, crustaceans, mollusks, algae and other organisms of value such as aquatic plants (e.g. Nelu ...
. Due to human introductions, it is now found in many tropical and subtropical habitats around the globe, where it can become an
invasive species An invasive species is an introduced species that harms its new environment. Invasive species adversely affect habitats and bioregions, causing ecological, environmental, and/or economic damage. The term can also be used for native spec ...
because of its robust nature. These same features make it a good species for aquaculture because it readily adapts to new situations.


Common names

The species is known by a number of
common name In biology, a common name of a taxon or organism (also known as a vernacular name, English name, colloquial name, country name, popular name, or farmer's name) is a name that is based on the normal language of everyday life; and is often con ...
s in places where they were introduced, including: * ''Blue kurper'' in
South Africa South Africa, officially the Republic of South Africa (RSA), is the Southern Africa, southernmost country in Africa. Its Provinces of South Africa, nine provinces are bounded to the south by of coastline that stretches along the Atlantic O ...
. * ''Mujair'' in Indonesia, the name derived from a Javanese breeder Moedjair. *''Daya'' in Pakistan * ''Jesus fish'' in the Elim area of Jamaica, named for their habit of multiplying * ''Tilapia negra'' ( lit. black tilapia) in
Colombia Colombia, officially the Republic of Colombia, is a country primarily located in South America with Insular region of Colombia, insular regions in North America. The Colombian mainland is bordered by the Caribbean Sea to the north, Venezuel ...


Description

The native,
wild-type The wild type (WT) is the phenotype of the typical form of a species as it occurs in nature. Originally, the wild type was conceptualized as a product of the standard "normal" allele at a locus, in contrast to that produced by a non-standard, " ...
Mozambique tilapia is laterally compressed, being deep bodied. It has long dorsal fins, the front part of which have
spine Spine or spinal may refer to: Science Biology * Spinal column, also known as the backbone * Dendritic spine, a small membranous protrusion from a neuron's dendrite * Thorns, spines, and prickles, needle-like structures in plants * Spine (zoology), ...
s. Native coloration is a dull greenish or yellowish, and weak banding may be seen. Adults reach up to in
standard length Fish measurement is the measuring of individual fish and various parts of fish anatomy, their anatomies, for data used in many areas of ichthyology, including Taxonomy (biology), taxonomy and fishery biology. Overall length Standard length (SL) is ...
and up to . It lives up to 11 years. Size and coloration may vary in captive and naturalized populations due to environmental and breeding pressures.


Hybridization

As with most species of tilapia, Mozambique tilapia have a high potential for hybridization. They are often crossbred with other tilapia species in aquaculture because purebred Mozambique tilapia don't grow as quickly and have a body shape poorly suited to cutting large
fillet Fillet may refer to: *Annulet (architecture), part of a column capital, also called a fillet *Fillet (aircraft), a fairing smoothing the airflow at a joint between two components *Fillet (clothing), a headband *Fillet (heraldry), diminutive of the ...
s; these hybrids may or may not be reported as hybrid fish, often being listed as the wild species, ''Oreochromis mossambicus''. Mozambique tilapia are hybridized because they naturally tolerate saltwater, a desirable trait in aquaculture, and the resultant offspring may inherit this trait. Also, hybrids between certain parent combinations (such as between Mozambique and
Wami tilapia The Wami tilapia is a tilapiine cichlid that grows to over 20 cm in length and is considered a useful food fish in Tanzania and the island of Zanzibar, which is recognized as a potential origin. It is tolerant of brackish water and grows we ...
) result in offspring that are predominantly or entirely male. Male tilapia are preferred in aquaculture as they grow faster and have a more uniform adult size than females. The "Florida Red" tilapia is a popular commercial hybrid of Mozambique and
blue tilapia The blue tilapia (''Oreochromis aureus'') is a species of tilapia, a fish in the family Cichlidae. Native to Northern and Western Africa, and the Middle East, through introductions it is now also established elsewhere, including parts of the Un ...
.


Distribution and habitat

The Mozambique tilapia is native to inland and coastal waters in southeastern Africa, from the
Zambezi The Zambezi (also spelled Zambeze and Zambesi) is the fourth-longest river in Africa, the longest east-flowing river in Africa and the largest flowing into the Indian Ocean from Africa. Its drainage basin covers , slightly less than half of t ...
basin in
Mozambique Mozambique, officially the Republic of Mozambique, is a country located in Southeast Africa bordered by the Indian Ocean to the east, Tanzania to the north, Malawi and Zambia to the northwest, Zimbabwe to the west, and Eswatini and South Afr ...
,
Malawi Malawi, officially the Republic of Malawi, is a landlocked country in Southeastern Africa. It is bordered by Zambia to the west, Tanzania to the north and northeast, and Mozambique to the east, south, and southwest. Malawi spans over and ...
,
Zambia Zambia, officially the Republic of Zambia, is a landlocked country at the crossroads of Central Africa, Central, Southern Africa, Southern and East Africa. It is typically referred to being in South-Central Africa or Southern Africa. It is bor ...
and
Zimbabwe file:Zimbabwe, relief map.jpg, upright=1.22, Zimbabwe, relief map Zimbabwe, officially the Republic of Zimbabwe, is a landlocked country in Southeast Africa, between the Zambezi and Limpopo Rivers, bordered by South Africa to the south, Bots ...
to Bushman River in South Africa's
Eastern Cape The Eastern Cape ( ; ) is one of the nine provinces of South Africa. Its capital is Bhisho, and its largest city is Gqeberha (Port Elizabeth). Due to its climate and nineteenth-century towns, it is a common location for tourists. It is also kno ...
province. It is threatened in its home range by the introduced
Nile tilapia The Nile tilapia (''Oreochromis niloticus'') is a species of tilapia, a cichlid occurring naturally in parts of Africa (such as its namesake Nile River) and the Levant, though numerous introduced populations exist outside its natural range. T ...
. In addition to competing for the same resources, the two readily hybridize. This has already been documented from the Zambezi and
Limpopo River The Limpopo River () rises in South Africa and flows generally eastward through Mozambique to the Indian Ocean. The term Limpopo is derived from Rivombo (Livombo/Lebombo), a group of Tsonga settlers led by Hosi Rivombo who settled in the mou ...
s, and it is expected that pure Mozambique tilapia eventually will disappear from both. Otherwise it is a remarkably robust and fecund fish, readily adapting to available food sources and breeding under suboptimal conditions. Among others, it occurs in rivers, streams, canals, ponds, lakes, swamps and
estuaries An estuary is a partially enclosed coastal body of brackish water with one or more rivers or streams flowing into it, and with a free connection to the open sea. Estuaries form a transition zone between river environments and maritime environm ...
, although it typically avoids fast-flowing waters, waters at high altitudes and the open sea. It inhabits waters that range from .


Invasiveness

The Mozambique tilapia or hybrids involving this species and other tilapia are invasive in many parts of the world outside their native range, having escaped from aquaculture or deliberately introduced to control mosquitoes. The Mozambique tilapia is listed by the Invasive Species Specialist Group as one of the top 100 worst invasive species in the world. It can harm native fish populations through competition for food and nesting space, as well as by directly consuming small fish. In Hawaii, striped mullet ''
Mugil cephalus The flathead grey mullet (''Mugil cephalus'') is an important food fish species in the mullet family Mugilidae. It is found in coastal temperate, tropical and subtropical waters worldwide. Its length is typically . It is known with numerous En ...
'' are threatened because of the introduction of this species. The population of hybrid Mozambique tilapia x
Wami tilapia The Wami tilapia is a tilapiine cichlid that grows to over 20 cm in length and is considered a useful food fish in Tanzania and the island of Zanzibar, which is recognized as a potential origin. It is tolerant of brackish water and grows we ...
in California's
Salton Sea The Salton Sea is a shallow, landlocked, highly salinity, saline endorheic lake in Riverside County, California, Riverside and Imperial County, California, Imperial counties in Southern California. It lies on the San Andreas Fault within the S ...
may also be responsible for the decline of the desert pupfish, '' Cyprinodon macularius''.


Behavior


Feeding

Mozambique tilapia are
omnivorous An omnivore () is an animal that regularly consumes significant quantities of both plant and animal matter. Obtaining energy and nutrients from plant and animal matter, omnivores digest carbohydrates, protein, fat, and fiber, and metabolize ...
. They can consume
detritus In biology, detritus ( or ) is organic matter made up of the decomposition, decomposing remains of organisms and plants, and also of feces. Detritus usually hosts communities of microorganisms that colonize and decomposition, decompose (Reminera ...
,
diatom A diatom (Neo-Latin ''diatoma'') is any member of a large group comprising several Genus, genera of algae, specifically microalgae, found in the oceans, waterways and soils of the world. Living diatoms make up a significant portion of Earth's B ...
s,
phytoplankton Phytoplankton () are the autotrophic (self-feeding) components of the plankton community and a key part of ocean and freshwater Aquatic ecosystem, ecosystems. The name comes from the Greek language, Greek words (), meaning 'plant', and (), mea ...
, invertebrates, small fry and vegetation ranging from
macroalgae Seaweed, or macroalgae, refers to thousands of species of Macroscopic scale, macroscopic, Multicellular organism, multicellular, ocean, marine algae. The term includes some types of ''Rhodophyta'' (red), ''Brown algae, Phaeophyta'' (brown) and ...
to rooted plants. This broad diet helps the species thrive in diverse locations. Due to their robust nature, Mozambique tilapias often over-colonize the habitat around them, eventually becoming the most abundant species in a particular area. When over-crowding happens and resources get scarce, adults will sometimes cannibalize the young for more nutrients. Mozambique tilapia, like other fish such as
Nile tilapia The Nile tilapia (''Oreochromis niloticus'') is a species of tilapia, a cichlid occurring naturally in parts of Africa (such as its namesake Nile River) and the Levant, though numerous introduced populations exist outside its natural range. T ...
and
trout Trout (: trout) is a generic common name for numerous species of carnivorous freshwater ray-finned fishes belonging to the genera '' Oncorhynchus'', ''Salmo'' and ''Salvelinus'', all of which are members of the subfamily Salmoninae in the ...
, are opportunistic
omnivores An omnivore () is an animal that regularly consumes significant quantities of both plant and animal matter. Obtaining energy and nutrients from plant and animal matter, omnivores digest carbohydrates, protein, fat, and fiber, and metabolize t ...
and will feed on
algae Algae ( , ; : alga ) is an informal term for any organisms of a large and diverse group of photosynthesis, photosynthetic organisms that are not plants, and includes species from multiple distinct clades. Such organisms range from unicellular ...
, plant matter, organic particles, small invertebrates and other fish. Feeding patterns vary depending on which food source is the most abundant and the most accessible at the time. In captivity, Mozambique tilapias have been known to learn how to feed themselves using demand feeders. During commercial feeding, the fish may energetically jump out of the water for food.


Social structure

Mozambique tilapias often travel in groups where a strict
dominance hierarchy In the zoological field of ethology, a dominance hierarchy (formerly and colloquially called a pecking order) is a type of social hierarchy that arises when members of animal social animal, social groups interact, creating a ranking system. Dif ...
is maintained. Positions within the hierarchy correlate with territoriality, courtship rate, nest size, aggression, and hormone production. In terms of social structure, Mozambique tilapias engage in a system known as lek-breeding, where males establish territories with dominance hierarchies while females travel between them. Social hierarchies typically develop because of competition for limited resources including food, territories, or mates. During the breeding season, males cluster around certain territory, forming a dense aggregation in shallow water. This aggregation forms the basis of the lek through which the females preferentially choose their mates. Reproductive success by males within the lek is highly correlated to social status and dominance. In experiments with captive tilapias, evidence demonstrates the formation of linear hierarchies where the alpha male participates in significantly more agonistic interactions. Thus, males that are higher ranked initiate much more aggressive acts than subordinate males. However, contrary to popular belief, Mozambique tilapias display more agonistic interactions towards fish that are farther apart in the hierarchy scale than they do towards individuals closer in rank. One hypothesis behind this action rests with the fact that aggressive actions are costly. In this context, members of this social system tend to avoid confrontations with neighboring ranks in order to conserve resources rather than engage in an unclear and risky fight. Instead, dominant individuals seek to bully subordinate tilapias both for an easy fight and to keep their rank.


Communication and aggression

The
urine Urine is a liquid by-product of metabolism in humans and many other animals. In placental mammals, urine flows from the Kidney (vertebrates), kidneys through the ureters to the urinary bladder and exits the urethra through the penile meatus (mal ...
of Mozambique tilapias, like many freshwater fish species, acts as a vector for communication amongst individuals.
Hormone A hormone (from the Ancient Greek, Greek participle , "setting in motion") is a class of cell signaling, signaling molecules in multicellular organisms that are sent to distant organs or tissues by complex biological processes to regulate physio ...
s and
pheromones A pheromone () is a secreted or excreted chemical factor that triggers a social response in members of the same species. Pheromones are chemicals capable of acting like hormones outside the body of the secreting individual, to affect the behavi ...
released with urine by the fish often affect the behavior and physiology of the opposite sex. Dominant males signal females through the use of a urinary odorant. Further studies have suggested that females respond to the ratio of chemicals within the urine, as opposed to the odor itself. Nevertheless, females are known to be able to distinguish between hierarchical rank and dominant vs. subordinate males through chemicals in urine. Urinary pheromones also play a part in male – male interaction for Mozambique tilapias. Studies have shown that male aggression is highly correlated with increased urination. Symmetrical aggression between males resulted in an increase in the release of urination frequency. Dominant males both store and release more potent urine during agonistic interactions. Thus, both the initial stage of lek formation and the maintenance of social hierarchy may highly depend on the males’ varying urinary output. Aggression amongst males usually involve a typical sequence of visual, acoustic, and tactile signals that eventually escalates to physical confrontation if no resolution is reached. Usually, conflict ends before physical aggression as fights are both costly and risky. Bodily damage may impede an individual's ability to find a mate in the future. In order to prevent cheating, in which individual may fake his own fitness, these aggressive rituals incur significant energetic costs. Thus, cheating is prevented by the sheer fact that the costs of initiating a ritual often outweigh the benefits of cheating. In this regard, differences between individuals in endurance plays a critical role in resolving the winner and the loser.


Reproduction

In the first step in the reproductive cycle for Mozambique tilapia, males excavate a nest into which a female can lay her eggs. After the eggs are laid, the male fertilizes them. Then the female stores the eggs in her mouth until the fry hatch; this act is called
mouthbrooding Mouthbrooding, also known as oral incubation and buccal incubation, is the care given by some groups of animals to their offspring by holding them in the mouth of the parent for extended periods of time. Although mouthbrooding is performed by a va ...
.Popma, 1999 One of the main reasons behind the aggressive actions of Mozambique tilapias is access to reproductive mates. The designation of Mozambique tilapias as an invasive species rests on their life-history traits: Tilapias exhibit high levels of parental care as well as the capacity to spawn multiple
broods Broods are a musical duo from Nelson, New Zealand, composed of Georgia Josiena Nott on lead vocals, with older brother and multi-instrumentalist Caleb Allan Joseph Nott on production and backing vocals. They released the single "Bridges (Broo ...
through an extended reproductive season, both contributing to their success in varying environments. In the lek system, males congregate and display themselves to attract females for matings. Thus, mating success is highly skewed towards dominant males, who tend to be larger, more aggressive, and more effective at defending territories. Dominant males also build larger nests for the
spawn Spawn or spawning may refer to: * Spawning, the eggs and sperm of aquatic animals Arts, entertainment and media * Spawn (character), a fictional character in the comic series of the same name and in the associated franchise ** ''Spawn: Armageddon' ...
. During courtship rituals, acoustic communication is widely used by the males to attract females. Studies have shown that females are attracted to dominant males who produce lower peak frequencies as well as higher pulse rates. At the end of mating, males guard the nest while females take both the eggs and the sperm into their mouth. Due to this, Mozambique tilapias can occupy many niches during spawning since the young can be transported in the mouth. These proficient reproductive strategies may be the cause behind their invasive tendencies. Male Mozambique tilapias synchronize breeding behavior in terms of
courtship Courtship is the period wherein some couples get to know each other prior to a possible marriage or committed romantic, ''de facto'' relationship. Courtship traditionally may begin after a betrothal and may conclude with the celebration of marri ...
activity and territoriality in order to take advantage of female spawning synchrony. One of the costs associated with this synchronization is the increase in competition among males, which are already high on the
dominance hierarchy In the zoological field of ethology, a dominance hierarchy (formerly and colloquially called a pecking order) is a type of social hierarchy that arises when members of animal social animal, social groups interact, creating a ranking system. Dif ...
. As a result, different mating tactics have evolved in these species. Males may
mimic In evolutionary biology, mimicry is an evolved resemblance between an organism and another object, often an organism of another species. Mimicry may evolve between different species, or between individuals of the same species. In the simples ...
females and sneak reproduction attempts when the dominant male is occupied. Likewise, another strategy for males is to exist as a floater, travelling between territories in an attempt to find a mate. Nevertheless, it is the dominant males who have the greatest reproductive advantage.


Parental care

Typically, Mozambique tilapias, like all species belonging to the genus ''
Oreochromis ''Oreochromis'' is a large genus of oreochromine cichlids, fishes endemic to Africa and the Middle East. A few species from this genus have been introduced far outside their native range and are important in aquaculture. Many others have very ...
'' and species like '' Astatotilapia burtoni'', are maternal mouthbrooders, meaning that spawn is incubated and raised in the mouth of the mother.
Parental care Parental care is a behavioural and evolutionary strategy adopted by some animals, involving a parental investment being made to the evolutionary fitness of offspring. Patterns of parental care are widespread and highly diverse across the animal k ...
is, therefore, almost exclusive to the female. Males do contribute by providing nests for the spawn before incubation, but the energy costs associated with nest production is low relative to mouthbrooding. Compared to nonmouthbrooders, both mouthbrooding and growing a new clutch of eggs is not energetically feasible. Thus, Mozambique tilapias arrest
oocyte An oocyte (, oöcyte, or ovocyte) is a female gametocyte or germ cell involved in reproduction. In other words, it is an immature ovum, or egg cell. An oocyte is produced in a female fetus in the ovary during female gametogenesis. The female ger ...
growth during
mouthbrooding Mouthbrooding, also known as oral incubation and buccal incubation, is the care given by some groups of animals to their offspring by holding them in the mouth of the parent for extended periods of time. Although mouthbrooding is performed by a va ...
to conserve energy. Even with oocyte arrest, females that mouthbrood take significant costs in body weight, energy, and low fitness. Hence, parental-offspring conflict is visible through the costs and benefits to the parents and the young. A mother caring for her offspring carries the cost of reducing her own individual fitness. Unlike most fish, Mozambique tilapias exhibit an extended maternal care period believed to allow social bonds to be formed.


Use in aquaculture

Mozambique tilapia are hardy, being easy to raise and harvest, making them a good aquacultural species. They have a mild, white flesh that is appealing to consumers. This species constitutes about 4% of the total tilapia aquaculture production worldwide, but is more commonly hybridized with other tilapia species. Tilapia are very susceptible to diseases such as whirling disease and ich. Mozambique tilapia are resistant to wide varieties of water quality issues and pollution levels. Because of these abilities they have been used as bioassay organisms to generate metal toxicity data for risk assessments of local freshwater species in
South Africa South Africa, officially the Republic of South Africa (RSA), is the Southern Africa, southernmost country in Africa. Its Provinces of South Africa, nine provinces are bounded to the south by of coastline that stretches along the Atlantic O ...
rivers. Mozambique tilapia were one of the species flown on the Bion-M No.1 spacecraft in 2013, but they all died due to equipment failure.


References


References

* * Courtenay W.R. Jr. 1989. Exotic fishes in the National Park System. Pages 237–252 in: Thomas L.K. (ed.). Proceedings of the 1986 conference on science in the national parks, volume 5. Management of exotic species in natural communities. U.S. National Park Service and George Wright Society, Washington, D.C. * Courtenay W.R. Jr., and C.R. Robins. 1989. Fish introductions: Good management, mismanagement, or no management? CRC Critical Reviews in Aquatic Sciences 1:159–172. * * Gupta M.V. and B.O. Acosta. 2004. A review of global tilapia farming practices. WorldFish Center P.O. Box 500 GPO, 10670, Penang, Malaysia. * * Moyle P.B. 1976. Inland fishes of California. University of California Press, Berkeley, CA. 330 p. * Popma, T. Tilapia Life History and Biology 1999 Southern Region Aquaculture Center * * Trewevas E. 1983. Tilapiine Fishes Of The Genera Sarotherodon, Oreochromis And Danakilia. British Museum Of Natural History, Publication Number 878.Comstock Publishing Associates. Ithaca, New York. 583 p. * Waal, Ben van der, 2002.
Another fish on its way to extinction?
'. Science in Africa.


External links


Photo of "Florida Red" hybrid
Retrieved 12 July 2007. Mozambique tilapia Freshwater fish of Africa Freshwater fish of South Africa Fish of Mozambique Fish of the Dominican Republic * Taxa named by Wilhelm Peters Fish described in 1852 Near threatened animals Vulnerable biota of Africa Space-flown life