العربية
中国
Français
Deutsch
Ελληνική
हिन्दी
Italiano
日本語
Português
Русский
EspañolCICHLID
'Cichlids' (IPA pronunciation: ) are fishes from the family 'Cichlidae' in the order Perciformes. The family Cichlidae, a major family of perciform fish, is both large and diverse. Estimates of the number of cichlid species range from 1,300 to 1,900, making it one of the three largest vertebrate families.[1][2] Cichlids span a wide range of body sizes, from species as small as 2.5 centimetres (1.0 in) in length (e.g. ''Neolamprologus multifasciatus'' ) to much larger species approaching 1 metre (3 ft) in length (e.g. ''Boulengerochromis'' and ''Cichla''). As a group, cichlids exhibit a similarly wide diversity of body shapes, ranging from strongly laterally compressed species (such as ''Altolamprologus'', ''Pterophyllum'', and ''Symphysodon'') through to species that are cylindrical and highly elongate (such as ''Julidochromis'', ''Teleogramma'', ''Teleocichla'', ''Crenicichla'', and ''Gobiocichla'').[3] Generally, however, cichlids tend to be of medium size, ovate in shape and slightly laterally compressed, and generally very similar to the North American sunfishes in terms of morphology, behaviour, and ecology.[4]
Some species, particularly the tilapiines are important food fishes, while others are valued game fish (eg. ''Cichla'' species). Many species, including the angelfish, oscars, and discus, are also highly valued in the aquarium trade.[5] Cichlids are also the family of vertebrates with by far the highest number of endangered species, most of these being from among the haplochromine group.[6] Cichlids are particularly well known for having evolved rapidly into a large number of closely related but morphologically diverse species within large lakes, particularly the African Rift Valley lakes of Tanganyika, and Victoria, and Malawi.[7][8] The diversity of cichlids in the African Great Lakes is important for the study of speciation in evolution. African Cichlid Fishes: Model Systems for Evolutionary Biology, Kornfield, Irv, , , Annual Review of Ecology and Systematics, 2000 Many cichlids that have been accidentally or deliberately released into freshwaters outside of their natural range have become nuisance species, for example tilapia in the southern United States.[9]
Anatomy and appearance
Relationships within the Labrodei.[10]
Cichlids are members of a group of perciform fish known as the Labroidei alongside the wrasses Labridae, damselfish Pomacentridae, and surfperches Embiotocidae. This very large grouping shares a single key trait: the fusion of the lower pharyngeal bones into a single tooth-bearing structure. A complex set of muscles allows the upper and lower pharyngeal bones to be used as a second set of jaws for processing food, allowing a division of labour between the "true jaws" (mandibles) and the "pharyngeal jaws". Cichlids in particular have evolved to be very efficient feeders that are able to capture and process a very wide variety of food items and this is assumed to be one reason why they are so diverse (see section on diet below). Cichlids have a great variability in body shape, ranging from compressed and disc-shaped (such as ''Symphysodon'') to elongate and cylindrical (such as ''Crenicichla''.[11]
The particular features of cichlids that distinguish them from the other Labroidei include:
★ A single nostril on each side of the forehead instead of two.
★ No bony shelf below the orbit of the eye.
★ The lateral line organ is divided into two sections, one on the upper half of the flank and a second along the midline of the flank from about halfway along the body to the base of the tail (except for genera ''Teleogramma'' and ''Gobiocichla'').
★ A distinctively shaped otolith.
★ The small intestine leaves the stomach from its left side, not from its right side as in other Labroidei.
Taxonomy
Kullander (1998) recognises eight subfamilies of cichlids: the Astronotinae, Cichlasomatinae, Cichlinae, Etroplinae, Geophaginae, Heterochromidinae, Pseudocrenilabrinae and Retroculinae. Nelson (2006), however, indicates that cichlid taxonomy is still greatly debated, and despite these attempts, classification of genera cannot yet be accurately given. A comprehensive system of assigning species to monophyletic genera is still lacking, and there is not complete agreement on what genera should be recognized in this family.
As an example of the extant problems in Cichlid taxonomy, Kullander published a phylogeny of the Cichlidae (details of which can be found here) i which the African genus ''Heterochromis'' ended up being placed phylogenetically within Neotropical (i.e., South American) Cichlds, although later papers arrived at different conclusions. Other extant problems (mentioned in the following paragraph) centre upon the identity of the putative common ancestor for the Lake Victoria superflock, and the precise ancestral lineages of the Tanganyikan Cichlids. However, the research is active and intensive (see below for just two of ''many'' recent papers), with new papers expected in the immediate future that may resolve some of the outstanding questions.
The paper ''Multilocus Phylogeny of Cichlid Fishes (Pisces: Perciformes): Evolutionary Comparison of Microsatellite and Single-Copy Nuclear Loci'' by Streelman, Zardoya, Meyer and Karl (1998) (Mol. Biol. Evol. 15(7):798–808. 1998, paper available as PDF here) provides comparisons between the phylogeny (due to Stiassny, 1991) as determined using morphological characteristics, and two maximum-pasrimony bootstrap consensus trees based upon analyses of two different gene loci, along with majority-rule trees and other similar phylogenetic trees based upon assorted gene locus analyses. There are some extant differences between the assorted trees at the genus level that require resolution, though the consensus from the various nuclear and mitochondrial DNA analyses in this and other papers is that the Cichlidae as a family is monophyletic. Further insights into the attractiveness of Cichlid taxonomy as a fertile area of research is given by the paper ''The species flocks of East African cichlid fishes: recent advances in molecular phylogenetics and population genetics'' by Salzburger and Meyer (Naturwissenschaften (2004) 91:277–290, paper available as PDF here), in which the advances made in the analysis of the phylogeny of the Lake Victoria superflock (among other East African Cichlids) is discussed in depth.
One problem that resulted in a major upheaval of Cichlid taxonomy was highlighted by Dr Humphrey Greenwood of the Natural History Museum, London, in a paper in 1977 (cited in TFH magazine, August 1977, with a follow up letter by Dr Greenwood in the November 1977 issue complaining about poor reportage of his work). Dentition (characteristics of tooth shape and arrangement) had been used prior to this date as classifying characteristics, which Greenwood demonstrated was invalid upon phylogenetic grounds because in many Cichlid species, dentition is an environmentally plastic characteristic (tooth shape changes in numerous species with age, due to wear etc), and cannot be relied upon universally across the family as a diagnostic characteristic upon which to base phylogenetic judgements. It is in the light of this that Cichlid taxonomy has undergone major revisions in the 30-plus years since, with the advent of genome sequencing and other technologies adding to the ever-growing body of data resulting in the still-fluid state of Cichlid taxonomy at the genus level.
Range and habitat
''Tilapia mariae'', caught on a hook and line, in Australia. Originally from Africa, the species established feral populations in Australia.[12]
Cichlids are the most species-rich non-Ostariophysan family in freshwaters worldwide. They are mainly freshwater fish that are most diverse in Africa and South America, with at least 900 species in the former and 291 in the latter. It is estimated that there will be about 1300 species in Africa alone when all are discovered and described. Substantial numbers are also found in Central America as far north as the Rio Grande in southern Texas, and Madagascar has its own distinctive fauna of cichlids phylogenetically only distantly related to those on the African mainland.[13] Endemic cichlids are largely absent in Asia except for four species in the Jordan Valley in the Middle East, one in Iran, and three in India and Sri Lanka. There are four species found in Cuba and Hispaniola. Europe, Australia, Antarctica, and most of North America do not have any native cichlids, although where environmental conditions are suitable, for example in Florida, Mexico, Japan and northern Australia, feral populations of cichlids have become established as exotics.[14][15][16][17][18][19]
Cichlids are largely freshwater fish and are less commonly found in brackish and salt water habitats, though many species will tolerate brackish water for extended periods; ''Cichlasoma urophthalmus'', for example, is equally at home in freshwater marshes and mangrove swamps, and can be found living and breeding in salt water environments such as the mangrove belts around barrier islands. Several species of tilapias (species of ''Tilapia'', ''Sarotherodon'', and ''Oreochromis'') are euryhaline and can disperse along some brackish coastlines between rivers. Only a few cichlids, however, are found primarily in brackish or salt water, most notably ''Etroplus maculatus'', ''Etroplus suratensis'', and ''Sarotherodon melanotheron''.[20]
Diet
The bumblebee cichlid, ''Pseudotropheus crabro'', is specialised in feeding on parasites from the catfish ''Bagrus meridionalis''[21]
Cichlids are astonishingly diverse in terms of diet. Many are primarily herbivores feeding on algae (e.g. ''Petrochromis'') and plants (e.g. ''Etroplus suratensis'') and small animals, particularly invertebrates, are only a small part of their diet. Some cichlids are detritivores and eat all types of organic material; among these species are the tilapiines of the genera ''Oreochromis'', ''Sarotherodon'', and ''Tilapia''.
Other cichlids are predatory and eat little if any plant matter. These include generalists that catch a variety of small animals including other fishes and insect larvae (e.g. ''Pterophyllum''), as well as variety of specialists. ''Trematocranus'' is a specialised snail-eater, while ''Pungu maclareni'' feeds on sponges. A number of cichlids feed on other fish, either whole or in part. ''Crenicichla'' are stealth-predators that lunge at small fish that pass by their hiding places, while ''Rhamphochromis'' are open water pursuit predators that chase down their prey. Rhamphochromis esox Oliver, M.K. Paedophagous cichlids such as the ''Caprichromis'' species eat other species' eggs or young (in some cases ramming the heads of mouthbrooding species to force them to disgorge their young).[22] The disputed feeding behavior of a paedophagous haplochromine cichlid (Pisces) observed and discussed, Wilhelm, W., , , Behaviour, 1980 [23][24] Among the more unusual feeding strategies are those of ''Corematodus'' spp., ''Docimodus evelynae'', ''Plecodus straeleni'' and all species of ''Perissodus'' , which feed on scales and fins of other fishes, a behaviour known as lepidophagy An example of "mimicry" in fishes, Trewavas, E., , , Nature, 1947 [25][26] along with the death mimicing behaviour of ''Nimbochromis'' and ''Parachromis'' species, which lay motionless, luring small fish to their side prior to ambush.[27][28]
Scientists believe it is this wide adaptability of feeding styles that has helped cichlids to inhabit such a wide range of habitats. It is largely the pharyngeal teeth (teeth in the throat) that allows the cichlids so many 'niche' feeding behaviours, i.e. the jaws may be used to hold or pick food, while the pharyngeal teeth are used to crush what was harvested.
Reproduction
A substrate brooding female managuense cichlid, ''Parachromis managuense'', guards a clutch of eggs in the aquarium.
Brood care
All species show some form of parental care for both eggs and larvae, often extended to free-swimming young until they are several weeks or months old. Species of this family have highly organized breeding activities.
Parental care falls into one of four categories:Substrate or open brooders, secretive cave brooders (also known as guarding speleophils[29]), and at least two types of mouthbrooding, ovophile mouthbrooding and larvophile mouthbrooding.[30]
Open or substrate brooding cichlids lay their eggs in the open on rocks, leaves or logs. Examples of open brooding cichlids include ''Pterophyllum'', ''Symphysodon'' spp and ''Anomalochromis thomasi''. In general, brooding biparental substrate brooding cichlids usually engage in differing roles with regard to protection and raising of the fry. Most commonly, the male parent patrols the pair's territory and repels intruders, while females more intensively tend the brood, fanning water over the eggs, removing infertile eggs and leading the school of fry while foraging. Despite this, both sexes are able to perform the full range of parenting behaviours. Secretive cave spawning cichlids lay their eggs in caves, crevices, holes or discarded mollusc shells (see also: shell brooding cichlids), frequently attaching the eggs to the roof of the chamber. Examples include ''Pelvicachromis'' spp., ''Archocentrus'' spp and ''Apistogramma'' spp. Communication between free-swimming fry and parents of both open and cave spawning cichlids has been observed for a number of cichlids in captivity and in the wild. Frequently this communication is based on body movements, such as shaking and pelvic fin flicking. In addition, parental substrate brooding cichlids assist in finding food resources for their fry. Parental behaviours such as leaf-turning, and fin-digging have been observed for a number of neotropical cichlid species.
Communal parental care, where multiple monogamous pairs care for a mixed school of young have also been observed for a number of cichlid species including: ''Amphilophus citrinellus'', ''Etroplus suratensis'' and ''Tilapia rendalli''.[31][32][33]
A female ''Cyphotilapia frontosa'' mouthbrooding fry which can be seen looking out her mouth
Ovophile mouthbrooders incubate their eggs in their mouths as soon as they are laid, and frequently continue to brood free-swimming fry in their mouths for several weeks. Examples of ovophile mouthbrooding cichlids include many of the cichlids endemic to the Rift Valley lakes (Lake Malawi, Lake Tanganyika and Lake Victoria) in east Africa eg: ''Maylandia'', ''Pseudotropheus'' and ''Tropheus'' along with some south american cichlids such as ''Geophagus steindachneri''. Larvophile mouthbrooding species lay the eggs in the open, or in a cave and upon hatching take the larvae into the mouth. Examples include some variants of ''Geophagus altifrons'', some ''Aequidens'', ''Gymnogeophagus'' and ''Satanoperca'' species. Regardless of whether eggs or larvae are subject to mouthbrooding, the vast majority of mouthbrooding cichlids are maternal mouthbrooders, that is the female mouthbroods the young. In the eretmodine cichlids (genera ''Spathodus'', ''Eretmodus'' and ''Tanganicodus''), some ''Sarotherodon'' species, ''Chromidotilapia guntheri'' and some ''Aequidens'' species, however, mouthbrooding is practiced by both the male and the female. Mysterious mouthbrooders, Coleman, R., , , Cichlid News, 1999 Paternal mouthbrooding, though rare, is also known to occur in the family eg. ''Sarotherodon melanotheron''.[34] This method is common and appears to have evolved independently in several groups of African cichlids.
Several cichlids, including discus (''Symphysodon'' spp.), some ''Amphilophus'' species, ''Etroplus'' and ''Uaru'' species are noted to feed their young with a skin secretion from mucous glands.
Mating system
Cichlids are either monogamous or polygamous in their mating system. The mating system of any given cichlid species is not consistently associated with the type of brood care the species employs. For example, although most monogamous cichlids are not mouthbrooding cichlids, ''Chromidotilapia'', ''Gymnogeophagus'', ''Spathodus'' and ''Tanganicodus'' are all monogamous mouthbrooders. In contrast, numerous open or cave spawning cichlids are polygamous, examples include ''Apistogramma'', ''Lamprologus'', ''Nannacara'' and ''Pelvicachromis''.[35]
''Lamprologus ocellatus'', a shell-dwelling cichlid, guarding fry
Endangered cichlids
According to the 2007 International Union for Conservation of Nature and Natural Resources red list 156 cichlid species are currently listed as vulnerable, 40 species are listed as endangered, while 69 species are listed as critically endangered. Six species, ''Haplochromis ishmaeli'', ''Haplochromis lividus'', ''Haplochromis perrieri'', ''Paretroplus menarambo'', ''Platytaeniodus degeni'' and ''Yssichromis'' sp. ''nov''. 'argens' are extinct in the wild, while at least 39 species, most from the genus ''Haplochromis'', have become extinct since the early 1990s.[36]
Lake Victoria
Because of the introduced Nile perch (''Lates niloticus'') and water hyacinth, deforestation causing siltation of water, and overfishing, many species of Lake Victoria cichlids have been wiped out or drastically reduced in the wild. By around 1980, fisheries of the lake yielded only 1 percent cichlids from all the catch, a drastic decline from 80 percent in the earlier years.
Cichlids as food and game fish
Although cichlids are mostly small and medium-sized fishes, a substantial number of species are notable as food and game fishes. With few thick rib bones and tasty flesh, artisan fishing of cichlids is not uncommon in Central America and South America, as well as areas surrounding the African rift lakes.[38] The most important food cichlids, however, are the tilapiines of North Africa. Fast growing, tolerant of stocking density, and highly adaptable, tilapiine species have been introduced and farmed extensively in many parts of Asia and are inceasingly common in other parts of the world. Production of farmed tilapia, at about 1.5 million tonnes annually with an estimated value of US$1.8 billion,[39] is about equal to that of salmon and trout. Unlike carnivorous salmon and trout, however, tilapia are mostly omnivorous and can feed on algae or any plant-based food. This reduces the cost of tilapia farming greatly and makes tilapia the ideal "aquatic chickens" of the trade.
In addition to being food fish, many large cichlids also make good game fish. The strong, hard-fighting peacock bass (''Cichla'' species) of South America is one of the most popular sportfish. It was intentionally released in many waters around the world. In Florida, this fish generates millions of hours of fishing and anglers' spendings of more than US$8 million a year.[40] Other cichlids preferred by anglers include the oscar, Mayan cichlid (''Cichlasoma urophthalmus''), and jaguar guapote (''Parachromis managuensis'').
Cichlids as aquarium fish
Although not particularly hardy, the discus, ''Symphysodon'' spp., has been popular among aquarium enthusiasts.
Since 1945, cichlids have become increasingly popular as aquarium fish[41][42][43][44][45][46] Cichlids are ideally suited as aquarium fish as many are small to medium-sized, easy to feed with a range of prepared fish foods, breed readily, and practice brood care, in captivity.
The most commonly encountered species in retail aquariums is ''Pterophyllum scalare'' from the Amazon River basin in tropical South America, known in the trade as the "angelfish" . Other popular or readily available species of cichlids include the oscar (''Astronotus ocellatus''), convict cichlid (''Archocentrus nigrofasciatus'') and discus (''Symphysodon'' spp.).
Species of cichlid can be kept in aquariums with other fish, however, many cichlids are predatory towards smaller fish. Conversely, some cichlids, such as ''Apistogramma'' or ''Julidochromis'' spp., can be timid in the aquarium. In such cases the use of dither fish is recommended.
Hybrid cichlids and selective breeding
The "red Texas" cichlid is not a Texas cichlid (''Herichthys cyanoguttatus'') but an intergeneric hybrid of ''Herichthys'' and ''Amphilophus'' parents.
Some cichlids have been found to hybridise with closely related species quite readily, both in the wild and under artificial conditions.[47] This is not particularly unusual, having been observed among other groups of fishes, such as European cyprinids.[48] What is unusual is the extent to which cichlid hybrids have been put to commercial use, in particular as food fish and as aquarium fish.[49] The red strain of tilapia hybrid, for example, is often preferred in aquaculture as they have faster growth rates. Tilapia hybridisation is also used to produce all-male populations to control stock density and prohibit reproduction in ponds.
The most ubiquitous aquarium hybrid is perhaps the blood parrot cichlid which is a cross of several species especially those from genus ''Amphilophus''. With a beak-shaped mouth, an abnormal spine, and an occasionally missing caudal fin (known as the "love heart" parrot cichlid), the fish has caused controversy among aquarium enthusiasts. Some has called blood parrot cichlid "the Frankenstein monster of the fish world."[50] Another notable hybrid, the flowerhorn cichlid, was very popular in some parts of Asia from 2001 until late 2003 and is believed to bring good luck to its owner.[51] The popularity of the flowerhorn cichlid declined in 2004,[52] resulting in many flowerhorn cichlids being released into the rivers and canals of Malaysia and Singapore where they pose a threat to endemic animal communities.[53]
A leucistic long-finned form of ''A. ocellatus''
Numerous cichlid species have also been the subject of selective breeding programmes to develop new ornamental strains for the aquarium trade. The most intensive selective breeding programs have involved angelfish and discus and many mutations that effect both colouration and finnage are known.[54][55] Many other cichlids have been selecively bred for albino, leucistic and xanthistic pigment mutations including oscars, convicts and ''Pelvicachromis pulcher''. Both dominant and recessively inherited pigment mutations have been observed for cichlids.[56] In convict cichlids, for example, a leucistic colouration is recessively inherited,[57] while in ''Oreochromis niloticus niloticus'' red colouration is caused by an dominantly inherited mutation.[58]
These efforts at selectively breeding may, however, have unintended consequences. For example, some selectively bred strains of ''Mikrogeophagus ramirezi'' have health and fertility problems.[59] Similarly, the inbreeding involved in selective breeding programmes can cause severe physical abnormalies such as the notched phenotype in angelfish.[60]
Genera
As of 2006, there were some 220 genera:
★ ''Acarichthys'' Eigenmann 1912 ★ ''Acaronia'' Myers 1940 ★ ''Aequidens'' Eigenmann & Bray 1894 ★ ''Alticorpus'' Stauffer & McKaye 1988 ★ ''Altolamprologus'' Poll 1986 ★ ''Amphilophus'' Agassiz, 1859 ★ ''Anomalochromis'' Greenwood 1985 ★ ''Apistogramma'' Regan 1913 ★ ''Apistogrammoides'' Meinken 1965 ★ ''Archocentrus'' Gill 1877 ★ ''Aristochromis'' Trewavas, 1935 ★ ''Astatoreochromis'' Pellegrin 1904 ★ ''Astatotilapia'' Pellegrin 1904 ★ ''Astronotus'' Swainson 1839 ★ ''Aulonocara'' Regan 1922 ★ ''Aulonocranus'' Regan 1920 ★ ''Australoheros'' Rican & Kullander 2006 ★ ''Baileychromis'' Poll 1986 ★ ''Bathybates'' Boulenger 1898 ★ ''Benitochromis'' Lamboj 2001 ★ ''Benthochromis'' Poll 1986 ★ ''Biotodoma'' Eigenmann & Kennedy 1903 ★ ''Biotoecus'' Eigenmann & Kennedy 1903 ★ ''Boulengerochromis'' Pellegrin 1904 ★ ''Buccochromis'' Eccles & Trewavas 1989 ★ ''Bujurquina'' Kullander 1986 ★ ''Callochromis'' Regan 1920 ★ ''Caprichromis'' Eccles & Trewavas 1989 ★ ''Caquetaia'' Fowler 1945 ★ ''Cardiopharynx'' Poll 1942 ★ ''Chaetobranchopsis'' Steindachner, 1875 ★ ''Chaetobranchus'' Heckel, 1840 ★ ''Chalinochromis'' Poll 1974 ★ ''Champsochromis'' Boulenger 1915 ★ ''Cheilochromis'' Eccles & Trewavas 1989 ★ ''Chetia'' Trewavas 1961 ★ ''Chilochromis'' Boulenger 1902 ★ ''Chilotilapia'' Boulenger 1908 ★ ''Chromidotilapia'' Boulenger 1898 ★ ''Cichla'' Bloch & Schneider 1801 ★ ''Cichlasoma'' Swainson 1839 ★ ''Cleithracara'' Kullander & Nijssen 1989 ★ ''Congochromis'' Stiassny & Schliewen 2007 ★ ''Copadichromis'' Eccles & Trewavas 1989 ★ ''Corematodus'' Boulenger 1897 ★ ''Crenicara'' Steindachner 1875 ★ ''Crenicichla'' Heckel 1840 ★ ''Ctenochromis'' Pfeffer 1893 ★ ''Ctenopharynx'' Eccles & Trewavas 1989 ★ ''Cunningtonia'' Boulenger 1906 ★ ''Cyathochromis'' Trewavas 1935 ★ ''Cyathopharynx'' Regan 1920 ★ ''Cyclopharynx'' Poll 1948 ★ ''Cynotilapia'' Regan 1922 ★ ''Cyphotilapia'' Regan 1920 ★ ''Cyprichromis'' Scheuermann 1977 ★ ''Cyrtocara'' Boulenger 1902 ★ ''Danakilia'' Thys van den Audenaerde 1969 ★ ''Dicrossus'' Steindachner 1875 ★ ''Dimidiochromis'' Eccles & Trewavas 1989 ★ ''Diplotaxodon'' Trewavas 1935 ★ ''Divandu'' Lamboj & Snoeks 2000 ★ ''Docimodus'' Boulenger 1897 ★ ''Eclectochromis'' Eccles & Trewavas 1989 ★ ''Ectodus'' Boulenger 1898 ★ ''Eretmodus'' Boulenger 1898 ★ ''Etia'' Schliewen & Stiassny 2003 ★ ''Etroplus'' Cuvier 1830 ★ ''Exochochromis'' Eccles & Trewavas 1989 ★ ''Fossorochromis'' Eccles & Trewavas 1989 ★ ''Genyochromis'' Trewavas 1935 ★ ''Geophagus'' Heckel 1840 ★ ''Gephyrochromis'' Boulenger 1901 ★ ''Gnathochromis'' Poll 1981 ★ ''Gobiocichla'' Kanazawa 1951 | ★ ''Grammatotria'' Boulenger 1899 ★ ''Greenwoodochromis'' Poll 1983 ★ ''Guianacara'' Kullander & Nijssen 1989 ★ ''Gymnogeophagus'' Miranda Ribeiro 1918 ★ ''Haplochromis'' Hilgendorf 1888 ★ ''Haplotaxodon'' Boulenger 1906 ★ ''Hemibates'' Regan 1920 ★ ''Hemichromis'' Peters 1857 ★ ''Hemitaeniochromis'' Eccles & Trewavas 1989 ★ ''Hemitilapia'' Boulenger 1902 ★ ''Herichthys'' Baird & Girard 1854 ★ ''Heros'' Heckel, 1840 ★ ''Herotilapia'' Pellegrin 1904 ★ ''Heterochromis'' Regan 1922 ★ ''Hoplarchus'' Kaup,1860 ★ ''Hoplotilapia'' Hilgendorf 1888 ★ ''Hypselecara'' Kullander 1986 ★ ''Hypsophrys'' Agassiz 1859 ★ ''Interochromis'' Yamaoka, Hori & Kuwamura 1988 ★ ''Iodotropheus'' Oliver & Loiselle 1972 ★ ''Iranocichla'' Coad 1982 ★ ''Ivanacara'' Romer & Hahn 2006 ★ ''Julidochromis'' Boulenger 1898 ★ ''Katria'' Stiassny & Sparks 2006 ★ ''Konia'' Trewavas 1972 ★ ''Krobia'' Kullander & Nijssen 1989 ★ ''Labeotropheus'' Ahl 1926 ★ ''Labidochromis'' Trewavas 1935 ★ ''Labrochromis'' Greenwood 1980 ★ ''Laetacara'' Kullander 1986 ★ ''Lamprologus'' Schilthuis 1891 ★ ''Lepidiolamprologus'' Pellegrin 1904 ★ ''Lestradea'' Poll 1943 ★ ''Lethrinops'' Regan 1922 ★ ''Lichnochromis'' Trewavas 1935 ★ ''Limbochromis'' Greenwood 1987 ★ ''Limnochromis'' Regan 1920 ★ ''Limnotilapia'' Regan 1920 ★ ''Lipochromis'' Greenwood 1980 ★ ''Lithochromis'' Lippitsch & Seehausen 1998 ★ ''Lobochilotes'' Boulenger 1915 ★ ''Macropleurodus'' Regan 1922 ★ ''Maylandia'' Meyer & Foerster 1984 ★ ★ (see also ''Metriaclima'') ★ ''Mazarunia'' Kullander 1990 ★ ''Mbipia'' Lippitsch & Seehausen 1998 ★ ''Mchenga'' Stauffer & Konings, 2006 ★ ''Melanochromis'' Trewavas, 1935 ★ ''Mesonauta'' Günther, 1862 ★ ''Microchromis'' Johnson 1975 ★ ''Mikrogeophagus'' Meulengracht-Madson 1968 ★ ''Myaka'' Trewavas 1972 ★ ''Mylacochromis'' Greenwood 1980 ★ ''Mylochromis'' Regan 1920 ★ ''Naevochromis'' Eccles & Trewavas 1989 ★ ''Nandopsis'' Gill, 1862 ★ ''Nannacara'' Regan]] 1905 ★ ''Nanochromis'' Pellegrin 1904 ★ ''Neetroplus'' Günther, 1867 ★ ''Neochromis'' Regan 1920 ★ ''Neolamprologus'' Colombe & Allgayer 1985 ★ ''Nimbochromis'' Eccles & Trewavas 1989 ★ ''Nyassachromis'' Eccles & Trewavas 1989 ★ ''Ophthalmotilapia'' Pellegrin 1904 ★ ''Oreochromis'' Günther 1889 ★ ''Orthochromis'' Greenwood 1954 ★ ''Otopharynx'' Regan 1920 ★ ''Oxylapia'' Kiener & Maugé 1966 ★ ''Pallidochromis'' Turner 1994 ★ ''Parachromis'' Agassiz 1859 ★ ''Paracyprichromis'' Poll 1986 ★ ''Paralabidochromis'' Greenwood 1956 ★ ''Parananochromis'' Greenwood 1987 ★ ''Paraneetroplus'' Regan 1905 | ★ ''Paratilapia'' Bleeker, 1868 ★ ''Paretroplus'' Bleeker, 1868 ★ ''Pelmatochromis'' Steindachner 1894 ★ ''Pelvicachromis'' Thys van den Audenaerde 1968 ★ ''Perissodus'' Boulenger 1898 ★ ''Petenia'' Günther, 1862 ★ ''Petrochromis'' Boulenger 1898 ★ ''Petrotilapia'' Trewavas 1935 ★ ''Pharyngochromis'' Greenwood 1979 ★ ''Placidochromis'' Eccles & Trewavas 1989 ★ ''Platytaeniodus'' Boulenger 1906 ★ ''Plecodus'' Boulenger 1898 ★ ''Prognathochromis'' Greenwood 1980 ★ ''Protomelas'' Eccles & Trewavas 1989 ★ ''Psammochromis'' Greenwood 1980 ★ ''Pseudocrenilabrus'' Fowler 1934 ★ ''Pseudosimochromis'' Nelissen 1977 ★ ''Pseudotropheus'' Regan 1922 ★ ''Pterochromis'' Trewavas 1973 ★ ''Pterophyllum'' Heckel 1840 ★ ''Ptychochromis'' Steindachner 1880 ★ ''Ptychochromoides'' Kiener & Mauge 1966 ★ ''Ptyochromis'' Greenwood 1980 ★ ''Pundamilia'' Seehausen & Lippitsch 1998 ★ ''Pungu'' Trewavas 1972 ★ ''Pyxichromis'' Greenwood 1980 ★ ''Reganochromis'' Whitley 1929 ★ ''Retroculus'' Eigenmann & Bray 1894 ★ ''Rhamphochromis'' Regan 1922 ★ ''Sargochromis'' Regan 1920 ★ ''Sarotherodon'' Rppell 1852 ★ ''Satanoperca'' Günther, 1862 ★ ''Schubotzia'' Boulenger 1914 ★ ''Schwetzochromis'' Poll 1948 ★ ''Sciaenochromis'' Eccles & Trewavas 1989 ★ ''Serranochromis'' Regan 1920 ★ ''Simochromis'' Boulenger 1898 ★ ''Spathodus'' Boulenger 1900 ★ ''Steatocranus'' Boulenger 1899 ★ ''Stigmatochromis'' Eccles & Trewavas 1989 ★ ''Stomatepia'' Trewavas 1962 ★ ''Symphysodon'' Heckel 1840 ★ ''Taeniacara'' Myers 1935 ★ ''Taeniochromis'' Eccles & Trewavas 1989 ★ ''Taeniolethrinops'' Eccles & Trewavas 1989 ★ ''Tahuantinsuyoa'' Kullander 1991 ★ ''Tangachromis'' Poll 1981 ★ ''Tanganicodus'' Poll 1950 ★ ''Teleocichla'' Kullander 1988 ★ ''Teleogramma'' Boulenger 1899 ★ ''Telmatochromis'' Boulenger 1898 ★ ''Theraps'' Günther, 1862 ★ ''Thoracochromis'' Greenwood 1979 ★ ''Thorichthys'' Meek 1904 ★ ''Thysochromis'' Daget 1988 ★ ''Tilapia'' Smith, 1840 See also: Tilapiine cichlids ★ ''Tomocichla'' Regan 1908 ★ ''Tramitichromis'' Eccles & Trewavas 1989 ★ ''Trematocara'' Boulenger 1899 ★ ''Trematocranus'' Trewavas 1935 ★ ''Triglachromis'' Poll & Thys van den Audenaerde 1974 ★ ''Tristramella'' Trewavas 1942 ★ ''Tridontochromis'' Greenwood 1980 ★ ''Tropheops'' Trewavas 1984 ★ ''Tropheus'' Boulenger 1898 ★ ''Tylochromis'' Regan 1920 ★ ''Tyrannochromis'' Eccles & Trewavas 1989 ★ ''Uaru'' Heckel 1840 ★ ''Variabilichromis'' Colombe & Allgayer 1985 ★ ''Vieja'' Fernandez-Yepez 1969 ★ ''Xenochromis'' Boulenger 1899 ★ ''Xenotilapia'' Boulenger 1899 ★ ''Xystichromis'' Greenwood 1980 |
Images of cichlids
:''Main gallery: ''
See also
★ Perciformes
★ Aquaculture
★ Fishkeeping
★ Aquarium
References
1.
2. Phylogeny and classification of neotropical fishes, , S.O., Kullander, EDIPUCRS, 1998,
3. The Cichlid Aquarium, , P.V., Loiselle, Tetra Press, 1994,
4. The Diversity of Fishes, Helfman G., Collette B., & Facey D., , , Blackwell Publishing, Inc., 1997,
5.
6. Captive breeding for the conservation of cichlid fishes, , G. M., Reid, Journal of Fish Biology, 1990
7. Out of Tanganyika: Genesis, explosive speciation, key-innovations and phylogeography of the haplochromine cichlid fishes, Salzburger W., Mack T., Verheyen E., Meyer A., , , BMC Evolutionary Biology, 2005
8. The cichlid diversity of Lake Malawi/Nyasa/Niassa: identification, distribution and taxonomy, Snoeks, J. (ed.), , , Cichlid Press, 2004,
9. Fact sheet for ''Oreochromis mossambicus'' (Peters, 1852) Gulf States Marine Fisheries Commission
10. Labroid intrarelationships revisited: morphological complexity, key innovations, and the study of comparative diversity, Stiassny, M.L.J., , , Bulletin of the Museum of Comparative Zoology, Harvard University, 1987
11. Fishes of the World, , Joseph, S., Nelson, John Wiley & Sons, Inc., 2006,
12. Priority management actions for alien freshwater fish species in Australia, Koehn, J.D., , , New Zealand Journal of Marine and Freshwater Research, 2004
13. Guide to Cichlids, , D. E., Boruchowitz, T.F.H. Publications, 2006, ISBN 0-7938-0584-8
14. Tilapia :: Far North Queensland ABC Far North Queensland
15. ''Archocentrus nigrofasciatus'', Convict cichlid Froese, R. and D. Pauly. Editors.
16. Hawai'i's native and exotic freshwater animals, Yamamoto, M.N., , , Mutual Publishing, 2000,
17. A field guide to freshwater fishes of North America north of Mexico, Page, L.M., , , Houghton Mifflin Company, 1991,
18. Fact Sheet for Tilapia zilli (Gervais, 1848) University of Southern Mississippi/College of Marine Sciences/Gulf Coast Research Laboratory
19. Nonindigenous Fishes of Florida - With a Focus on South Florida
20. Brackish-Water Fishes, Frank Schäfer, , , Aqualog, 2005, ISBN 3-936027-82-X (English), ISBN 3-936027-81-1 (German)
21. ''Melanochromis crabro'' sp. nov.: a cichlid fish from Lake Malawi which feeds on ectoparasites and catfish eggs, Ribbink, A.J., , , Netherlands Journal of Zoology, 1982 .
22. Head ramming behaviour by three paedophagous cichlids in Lake Malawi, Africa, McKaye, K.R., , , Animal Behaviour, 1983
23. Paedophagia among cichlid fishes of Lake Victoria and Lake Malawi/Nyasa, Ribbink, A.J., , , South African Journal of Science, 1997
24. Paedophagy in Malawi cichlids, Konings, A., , , Cichlid News, 2007
25. A taxonomic study of the genus ''Docimodus'' Boulenger (Pisces, Cichlidae) a group of fishes with unusual feeding habits from Lake Malawi, Eccles, D.H., , , Zoological Journal of the Linnean Society, 1976
26. Occasional egg-eating by the scale-eater ''Plecodus straeleni'' (Cichlidae) of Lake Tanganyika, Nshombo, M., , , Environmental Biology of Fishes, 1991
27. Feigning death in the Central American cichlid ''Parachromis friedrichsthalii'', Tobler, M., , , Journal of Fish Biology, 2005
28. Field observation on death feigning: a unique hunting behavior by the predatory cichlid, ''Haplochromis livingstoni'', of Lake Malawi, McKaye, K.R., , , Environmental Biology of Fishes, 1981
29. Reproductive guilds of fishes: a proposal. and definition, Balon, E.K., , , Journal of the Fisheries Research Board of Canada, 1975
30. Cichlid Fishes: behaviour, ecology and evolution, Keenleyside, M.H.A., , , Chapman and Hall, 1991,
31. Communal Care and Kidnapping of Young by Parental Cichlids, McKaye, K.R., , , Evolution, 1977
32. Ethology and ecology of cichlid fishes of the genus ''Etroplus'' in Sri Lanka: preliminary findings, Ward, J.A., , , Environmental Biology of Fishes, 1977
33. Nest-building and communal care of young by ''Tilapia rendalli dumeril'' (pisces, cichlidae) in Lake Malawi, Ribbink, A.J., , , Environmental Biology of Fishes, 1981
34. Paternal Mouthbrooding in the Black-Chinned Tilapia, ''Sarotherodon melanotheron'' (Pisces: Cichlidae): Changes in Gonadal Steroids and Potential for Vitellogenin Transfer to Larvae, Kishida, M., , , Hormones and Behavior, 2000
35. Alternative male mating acttics in a cichlid, ''Pelvicachromis pulcher'': a comparison of reproductive effort and success, Martin, E., , , Behavioral Ecology and Sociobiology, 1997
36. IUCN 2006. 2006 IUCN Red List of Threatened Species.
38. The Cichlid Fishes, , G. W., Barlow, Cambridge, MA: Perseus Publishing, 2000, ISBN 0-7382-0376-9
39. De Silva, S.S; Subasinghe, R.P.; Bartley, D.M.; Lowther, A. :Tilapias as Alien Aquatics in Asia and the Pacific: A Review. ''FAO Fisheries Technical Paper.'' No. 453, 2004. [1]
40. Fact Exotic Freshwater Fishes Florida Fish and Wildlife Conservation Commission
41. Keenleyside MHA (1991) ''Cichlid Fishes: behaviour, ecology and evolution'' Chapman and Hall, London.
42. Aquarium Atlas, , Rüdiger. Editor., Riehl, Tetra Press, 1996. 5th Edn.,
43. Sands D (1994) ''A fishkeepers guide to Central American cichlids.'' Tetra Press. Belgium pg 59-60.
44. Mills D (1993) ''Aquarium Fish'' Harper Collins ISBN 0-7322-5012-9
45. Konings A (1997) ''Back to nature guide to Malawi Cichlids'' Druckhaus Beltz, Germany. p. 13-23
46. Leibel WS (1993) ''A fishkeepers guide to South American cichlids.'' Tetra Press. Belgium pg 12-14.
47. Smith, P. F., Konings, A., and Kornfield I.: Hybrid origin of a cichlid population in Lake Malawi: implications for genetic variation and species diversity. ''Molecular Ecology'' '12', pp 2497–2504, 2003 [2]
48. Wood, A. B., and Jordan, D. R.: Fertility of roach × bream hybrids, ''Rutilus rutilus'' (L.) × ''Abramis brama'' (L.), and their identification. ''Journal of Fish Biology'' '30', pp 249-261, 1987
49. Frequently asked questions on Parrot cichlids Matt Clarke
50. It's The Frankenstein Monster Of The Fish World: The Blood Parrot!
51. Arnold W (2003) Singapore's 'lucky' pet Luohan can outnumber people in homes. International Herald Tribune. July 1.
52. ''Crayfish the latest fad among pet lovers'' New Straits Times (Malaysia) (2004) 3rd September
53. ''Flower Horn: Joy in homes, a pest in rivers''. New Straits Times (Malaysia) (2004) 14th July.
54. Norton J (1982) Angelfish genetics ''Freshwater And Marine Aquarium magazine'' '5:'(4)
55. Koh TL, Khoo G, Fan LQ, Phang VPE (1999) Genetic diversity among wild forms and cultivated varieties of Discus (Symphysodon spp.) as revealed by Random Amplified Polymorphic DNA (RAPD) fingerprinting ''Aquaculture'' '173:'485-497.
56. Kornfield I (1991) Genetics. In: ''Cichlid Fishes: behaviour, ecology and evolution'' Ed. Keenleyside MHA. Chapman and Hall, London. p. 109-115.
57. Itzkovich J, Rothbard S, Hulata G (1981) Inheritance of pink body colouration in cichlasoma nigrofasciatum Günther (Pisces, Cichlidae). ''Genetica'' '55:' 15-16.
58. McAndrew CJ, Roubal FR, Roberts RJ, Bullock AM, McEwan IM (1988) The genetics and history of red, blond, and associated color variants in ''Oreochromis niloticus'' ''Genetica'' '76:'172.
59. Linke H, Staeck L (1994) ''American cichlids I: Dwarf Cichlids. A handbook for their identification, care and breeding.'' Tetra Press. Germany. ISBN 1-56465-168-1
60. Norton J (1994) Notched - An Angelfish Deformity ''Freshwater And Marine Aquarium magazine'' '17:'(3)
Further reading
★ Barlow, G. W. (2000). ''The Cichlid fishes''. Cambridge MA: Perseus Publishing.
★ : National Museum of Natural History, Washington, D.C., 2004-05-11).
External links
★ Lost African Lake Spawned Fish Diversity "Beyond Belief" Discussing the biodiversity of cichlids.
★ FishBase.org The most current listing of cichlid taxonomy
★ The Cichlid Fishes of Lake Malawi by Dr. Michael Oliver.
★ An incomplete listing of cichlid genera and species, with phylogenetic context
This article provided by Wikipedia. To edit the contents of this article, click here for original source.
psst.. try this: add to faves
Featured Companies
| Golf Holidays International |
