Mozambique tilapia

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Mozambique tilapia
Oreochromis mossambicus.JPG
Scientific classification OOjs UI icon edit-ltr.svg
Domain: Eukaryota
Kingdom: Animalia
Phylum: Chordata
Class: Actinopterygii
Order: Cichliformes
Family: Cichlidae
Genus: Oreochromis
Species:
O. mossambicus
Binomial name
Oreochromis mossambicus
Synonyms
  • Chromis mossambicusW. K. H. Peters, 1852
  • Chromis niloticus var. mossambicusW. K. H. Peters, 1852
  • Sarotherodon mossambicus(W. K. H. Peters, 1852)
  • Tilapia mossambica(W. K. H. Peters, 1852)
  • Tilapia mossambica mossambica(W. K. H. Peters, 1852)
  • Tilapia mossambicus(W. K. H. Peters, 1852)
  • Chromis dumerilii Steindachner, 1864
  • Tilapia dumerilii(Steindachner, 1864)
  • Chromis vorax Pfeffer, 1893
  • Tilapia vorax(Pfeffer, 1893)
  • Chromis natalensis M. C. W. Weber, 1897
  • Sarotherodon mossambicus natalensis(M. C. W. Weber, 1897)
  • Tilapia natalensis(M. C. W. Weber, 1897)
  • Tilapia arnoldi Gilchrist & W. W. Thompson, 1917
  • Oreochromis mossambicus bassamkhalafiKhalaf, 2009

The Mozambique tilapia (Oreochromis mossambicus) is an oreochromine cichlid 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. Due to human introductions, it is now found in many tropical and subtropical habitats around the globe, where it can become an invasive species because of its robust nature. These same features make it a good species for aquaculture because it readily adapts to new situations. It is known as black tilapia in Colombia [2] and as blue kurper in South Africa. [3]

Contents

Description

The native Mozambique tilapia is laterally compressed, and has a deep body with long dorsal fins, the front part of which have spines. Native coloration is a dull greenish or yellowish, and weak banding may be seen. Adults reach up to 39 cm (15 in) in standard length and up to 1.1 kg (2.4 lb). [4] Size and coloration may vary in captive and naturalized populations due to environmental and breeding pressures. It lives up to 11 years. [4]

Distribution and habitat

An adult male in breeding condition Oreochromis mossambicus.jpg
An adult male in breeding condition

The Mozambique tilapia is native to inland and coastal waters in southeastern Africa, from the Zambezi basin in Mozambique, Malawi, Zambia and Zimbabwe to Bushman River in South Africa's Eastern Cape province. [1] [5] It is threatened in its home range by the introduced Nile tilapia. In addition to competing for the same resources, the two readily hybridize. [1] [6] This has already been documented from the Zambezi and Limpopo Rivers, and it is expected that pure Mozambique tilapia eventually will disappear from both. [1]

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, although it typically avoids fast-flowing waters, waters at high altitudes and the open sea. [1] [4] It inhabits waters that range from 17 to 35 °C (63–95 °F). [4] [7]

Invasiveness

Mozambique tilapia from West Bengal, India. Mozambique tilapia.jpg
Mozambique tilapia from West Bengal, India.

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 been deliberately introduced to control mosquitoes. [8] The Mozambique tilapia has been nominated by the Invasive Species Specialist Group as one of the 100 worst invasive species in the world. [9] It can harm native fish populations through competition for food and nesting space, as well as by directly consuming small fish. [10] In Hawaii, striped mullet Mugil cephalus are threatened because of the introduction of this species. The population of hybrid Mozambique tilapia x Wami tilapia in California's Salton Sea may also be responsible for the decline of the desert pupfish, Cyprinodon macularius . [11] [12] [13]

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 grow slowly and have a body shape poorly suited to cutting large fillets. However, Mozambique tilapia have the desirable trait of being especially tolerant of salty water. [14] Also, hybrids between certain parent combinations (such as between Mozambique and Wami tilapia) result in offspring that are all or predominantly 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. [15]

Behavior

Feeding

Mozambique tilapia are omnivorous. They can consume detritus, diatoms, phytoplankton, [16] invertebrates, small fry and vegetation ranging from macroalgae to rooted plants. [17] [18] 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 and trout, are opportunistic omnivores and will feed on algae, plant matter, organic particles, small invertebrates and other fish. [19] 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. [20]

Social structure

Mozambique tilapias often travel in groups where a strict dominance hierarchy is maintained. Positions within the hierarchy correlate with territoriality, courtship rate, nest size, aggression, and hormone production. [21] 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. [22] 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. [23]

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. [24]

Communication and aggression

Urine in Mozambique tilapias, like many freshwater fish species, acts as a vector for communication amongst individuals. Hormones and pheromones 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. [25]

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. [26]

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. [27] 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 through an extended reproductive season, both contributing to their success in varying environments. [28] 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. [22] 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. [29] These proficient reproductive strategies may be the cause behind their invasive tendencies.

Male Mozambique tilapias synchronize breeding behavior in terms of courtship 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. As a result, different mating tactics have evolved in these species. Males may mimic 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. [30]

Parental care

Typically, Mozambique tilapias, like all species belonging to the genus Oreochromis and species like Astatotilapia burtoni , are maternal mouthbrooders, meaning that spawn is incubated and raised in the mouth of the mother. Parental care 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 growth during mouthbrooding to conserve energy. [31] 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. [32]

Use in aquaculture

An albino strain has been developed in captivity TilapiamossambicaBCN.jpg
An albino strain has been developed in captivity

Mozambique tilapia are hardy individuals that are 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. [33] Tilapia are very susceptible to diseases such as whirling disease and ich. [27] 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 rivers. [34]

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. [35]

Other names

The species is known by a number of other names including:

Related Research Articles

<span class="mw-page-title-main">Cichlid</span> Family of fishes

Cichlids are fish from the family Cichlidae in the order Cichliformes. Traditionally Cichlids were classed in a suborder, the Labroidei, along with the wrasses (Labridae), in the order Perciformes, but molecular studies have contradicted this grouping. On the basis of fossil evidence, it first appeared in Tanzania during the Eocene epoch, about 46–45 million years ago; however, molecular clock estimates have placed the family's origin as far back as 67 million years ago, during the late Cretaceous period. The closest living relative of cichlids is probably the convict blenny, and both families are classified in the 5th edition of Fishes of the World as the two families in the Cichliformes, part of the subseries Ovalentaria. This family is large, diverse, and widely dispersed. At least 1,650 species have been scientifically described, making it one of the largest vertebrate families. New species are discovered annually, and many species remain undescribed. The actual number of species is therefore unknown, with estimates varying between 2,000 and 3,000.

<span class="mw-page-title-main">Tilapia</span> Common name for many species of fish

Tilapia is the common name for nearly a hundred species of cichlid fish from the coelotilapine, coptodonine, heterotilapine, oreochromine, pelmatolapiine, and tilapiine tribes, with the economically most important species placed in the Coptodonini and Oreochromini. Tilapia are mainly freshwater fish inhabiting shallow streams, ponds, rivers, and lakes, and less commonly found living in brackish water. Historically, they have been of major importance in artisanal fishing in Africa, and they are of increasing importance in aquaculture and aquaponics. Tilapia can become a problematic invasive species in new warm-water habitats such as Australia, whether deliberately or accidentally introduced, but generally not in temperate climates due to their inability to survive in cold water.

<span class="mw-page-title-main">Mouthbrooder</span> Animal that cares for its offspring by holding them its mouth

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 variety of different animals, such as the Darwin's frog, fish are by far the most diverse mouthbrooders. Mouthbrooding has evolved independently in several different families of fish.

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, where it may have been introduced by man. It is tolerant of brackish water and grows well in saline pools, making it particularly suitable for aquaculture by communities living close to the sea. Like other tilapia it is an omnivore and will feed on algae, plants, small invertebrates, and detritus. The common name refers to the Wami River.

<i>Tropheus moorii</i> Species of fish

Tropheus moorii is a species of cichlid endemic to Lake Tanganyika in Africa. Over 40 different color morphs of this species are dispersed throughout the lake, ranging from dark green to flame red and yellow. They mostly feed on filamentous algae on the rocky shallows they inhabit. T. moorii is a maternal mouthbrooder, so eggs are fertilized and young are carried in the mouth of the female while they hatch and develop.

<span class="mw-page-title-main">Nile tilapia</span> Species of fish

The Nile tilapia is a species of tilapia, a cichlid fish native to parts of Africa and the Levant, particularly Israel and Lebanon. Numerous introduced populations exist outside its natural range. It is also commercially known as mango fish, nilotica, or boulti.

<i>Oreochromis</i> Genus of fishes

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 small ranges; some are seriously threatened, and O. ismailiaensis and O. lidole possibly are extinct. Although Oreochromis primarily are freshwater fish of rivers, lakes and similar habitats, several species can also thrive in brackish waters and some even survive in hypersaline conditions with a salinity that far surpasses that of seawater. In addition to overfishing and habitat loss, some of the more localized species are threatened by the introduction of other, more widespread Oreochromis species into their ranges. This is because they—in addition to competing for the local resources—often are able to hybridize.

<span class="mw-page-title-main">Banggai cardinalfish</span> Species of fish

The Banggai cardinalfish is a small tropical cardinalfish in the family Apogonidae. It is the only member of its genus. This attractive fish is popular in the aquarium trade. It is among the relatively few marine fish to have been bred regularly in captivity, but significant numbers are still captured in the wild and it is now an endangered species. The detrimental impact of humans on its environment and certain fatal diseases threaten this species' numbers significantly. Iridovirus diseases are known to be significant reason for fish mortality.

<span class="mw-page-title-main">Butterfly splitfin</span> Species of fish

The butterfly splitfin or butterfly goodeid is a bony fish from the monotypic genus Ameca of the splitfin family (Goodeidae). It was formerly found throughout the Ameca River drainage in Mexico; the type locality is Rio Teuchitlán in the vicinity of Teuchitlán, Jalisco. The species was only ever found in an area about 10 miles (15 km) in diameter.

<span class="mw-page-title-main">Aquaculture of tilapia</span> Third most important fish in aquaculture after carp and salmon

Tilapia has become the third most important fish in aquaculture after carp and salmon; worldwide production exceeded 1.5 million metric tons in 2002 and increases annually. Because of their high protein content, large size, rapid growth, and palatability, a number of coptodonine and oreochromine cichlids—specifically, various species of Coptodon, Oreochromis, and Sarotherodon—are the focus of major aquaculture efforts.

Throughout much of the tropics, tilapiine cichlids native to Africa and the Levant have been widely introduced into a variety of aquatic systems. In the U.S. states of Florida and Texas, tilapia were originally introduced to curtail invasive plants. In an effort to meet the growing demand for tilapia, humans have farmed these fish in countries around the world. Capable of establishing themselves into new ponds and waterways, many tilapia have escaped aquaculture facilities across much of Asia, Africa, and South America. In other cases, tilapia have been established into new aquatic habitats via aquarists or ornamental fish farmers.

<i>Astatotilapia burtoni</i> Species of fish

Astatotilapia burtoni is a species of fish in the family Cichlidae.

<i>Oreochromis leucostictus</i> Species of fish

Oreochromis leucostictus is a species of cichlid native to Albertine Rift Valley lakes and associated rivers in DR Congo and Uganda. It has now been introduced widely elsewhere East Africa, and is believed to have negative ecological impact, particularly on native tilapias. This species is reported to reach a standard length of up to 36.3 cm (14.3 in), but is usually much smaller. It is exploited by small-scale fishery and aquaculture operations.

<i>Oreochromis lidole</i> Species of fish

Oreochromis lidole is a species of freshwater fish in the family Cichlidae. This tilapia is native to Malawi, Mozambique and Tanzania, where it is found in Lake Malawi, Lake Malombe, the Shire River and perhaps some crater lakes further north. It is important in fisheries, but has drastically declined; it may already be extinct. This oreochromine cichlid is locally called chambo, a name also used for two other closely related species found in the same region, O. karongae and O. squamipinnis.

<i>Oreochromis variabilis</i> Species of fish

Oreochromis variabilis, the Victoria tilapia, is a species of African cichlid native to Lake Victoria and its tributaries, Lake Kyoga, Lake Kwania, and Lake Bisina (Salisbury), as well as being found in the Victoria Nile above Murchison Falls. This species can reach a standard length of 30 cm (12 in). This species is important to local commercial fisheries and is potentially important in aquaculture. It is also found in the aquarium trade.

<span class="mw-page-title-main">Mango tilapia</span> Species of fish

The mango tilapia is a species of fish from the cichlid family that is native to fresh and brackish waters in Africa and the Levant. Other common names include Galilaea tilapia, Galilean comb, Galilee St. Peter's fish, and St. Peter's fish. This is a relatively large cichlid at up to 41 centimetres (16 in) in total length and about 1.6 kilograms (3.5 lb) in weight. It is very important to local fisheries and the species is also aquacultured.

<span class="mw-page-title-main">Redbelly tilapia</span> Species of fish

The redbelly tilapia, also known as the Zille's redbreast tilapia or St. Peter's fish, is a species of fish in the cichlid family. This fish is found widely in fresh and brackish waters in the northern half of Africa and the Middle East. Elsewhere in Africa, Asia, Australia and North America, it has been introduced as a food fish or as a control of aquatic vegetation. Where introduced, it sometimes becomes invasive, threatening the local ecology and species. The redbelly tilapia is an important food fish and sometimes aquacultured.

<span class="mw-page-title-main">Oreochromini</span> Tribe of fishes

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 Tilapiini but more recent workers have found that the Tilapiini sensu lato is paraphyletic. Despite this change, species in Oreochromini are still referred to by the common name tilapia and some of the most important tilapia in aquaculture —certain species of Oreochromis and Sarotherodon— are part of this tribe. In contrast, several species have small ranges and are seriously threatened; a few are already extinct or possibly extinct.

<i>Oreochromis mortimeri</i> Species of fish

Oreochromis mortimeri, the Kariba tilapia or kurper bream, is a species of cichlid, formerly classified as a Tilapiine cichlid but now placed in the genus Oreochromis, the type genus of the tribe Oreochromini of the subfamily Pseudocrenilabrinae. It is found in the rivers of south central Africa especially the middle Zambezi where it is endangered by the spread of invasive congener Oreochromis niloticus.

References

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  2. ↑ "Listado oficial de especies invasoras para Colombia | Parques Nacionales Naturales de Colombia".
  3. ↑ "Big Bass". Archived from the original on 2019-06-14. Retrieved 2012-03-22.
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  7. ↑ Ford, A.G.P.; et al. (2019). "Molecular phylogeny of Oreochromis (Cichlidae: Oreochromini) reveals mito-nuclear discordance and multiple colonisation of adverse aquatic environments" (PDF). Mol. Phylogenet. Evol. 136: 215–226. doi:10.1016/j.ympev.2019.04.008. PMID   30974200. S2CID   109938635.
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  9. ↑ Courtenay 1989
  10. ↑ Courtenay et al. 1974
  11. ↑ Courtenay and Robins 1989
  12. ↑ Swift et al. 1993
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  15. ↑ "Culture of Hybrid Tilapia: A Reference Profile". 2018-11-21.
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  17. ↑ Mook 1983
  18. ↑ Trewevas 1983
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  21. ↑ Oliveira, Rui F.; Vitor C. Almada; Adelino V. M. Canario (1996). "Social Modulation of Sex Steroid Concentrations in the Urine of Male Cichlid Fish Oreochromis mossambicus". Hormones and Behavior. 30 (1): 2–12. doi:10.1006/hbeh.1996.0002. hdl: 10400.1/3206 . PMID   8724173. S2CID   1951640.
  22. 1 2 Amorim, M. Clara P.; Almada, Vitor C. (1 March 2005). "The outcome of male–male encounters affects subsequent sound production during courtship in the cichlid fish Oreochromis mossambicus". Animal Behaviour. 69 (3): 595–601. doi:10.1016/j.anbehav.2004.06.016. hdl: 10400.12/1440 . S2CID   34065315.
  23. ↑ Barata, Eduardo N.; Fine, Jared M.; Hubbard, Peter C.; Almeida, Olinda G.; Frade, Pedro; Sorensen, Peter W.; Canário, Adelino V. M. (1 April 2008). "A Sterol-Like Odorant in the Urine of Mozambique Tilapia Males Likely Signals Social Dominance to Females". Journal of Chemical Ecology. 34 (4): 438–449. doi:10.1007/s10886-008-9458-7. PMID   18379847. S2CID   33463442.
  24. ↑ Oliveira, R.F.; V.C. Almada (1996). "Dominance hierarchies and social structure in captive groups of the Mozambique tilapia Oreochromis mossambicus (Teleostei Cichlidae)". Ethology Ecology & Evolution. 8: 39–55. doi:10.1080/08927014.1996.9522934. hdl: 10400.12/1330 .
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  27. 1 2 Popma, 1999
  28. ↑ Russell, D. J.; Thuesen, P. A.; Thomson, F. E. (1 May 2012). "Reproductive strategies of two invasive tilapia species Oreochromis mossambicus and Tilapia mariae in northern Australia". Journal of Fish Biology. 80 (6): 2176–2197. Bibcode:2012JFBio..80.2176R. doi:10.1111/j.1095-8649.2012.03267.x. PMID   22551176.
  29. ↑ Amorim, M. C. P.; Fonseca, P. J.; Almada, V. C. (1 March 2003). "Sound production during courtship and spawning of Oreochromis mossambicus: male-female and male-male interactions" (PDF). Journal of Fish Biology. 62 (3): 658–672. Bibcode:2003JFBio..62..658A. doi:10.1046/j.1095-8649.2003.00054.x. hdl: 10400.12/1452 . S2CID   28236698. Archived from the original (PDF) on 4 February 2021.
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  32. ↑ Russock, Howard I. (March 1986). "Preferential Behaviour of Sarotherodon (Oreochromis) mossambicus (Pisces: Cichlidae) Fry to Maternal Models and Its Relevance to the Concept of Imprinting". Behaviour. 96 (3/4): 304–321. doi:10.1163/156853986x00531.
  33. ↑ Gupta and Acosta 2004
  34. ↑ Mashifane, TB; Moyo, NAG (29 October 2014). "Acute toxicity of selected heavy metals to Oreochromis mossambicus fry and fingerlings". African Journal of Aquatic Science. 39 (3): 279–285. doi:10.2989/16085914.2014.960358. S2CID   85396607.
  35. ↑ Zak, Anatoly. "Bion (12KSM) satellite". RussianSpaceWeb.com. Archived from the original on 9 June 2013. Retrieved 20 May 2013.

References

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