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Cichlid

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Cichlid
Temporal range:48.6–0Ma[1][2]Early Eocene-present
Ambuna
Scientific classificationEdit this classification
Domain: Eukaryota
Kingdom: Animalia
Phylum: Chordata
Class: Actinopterygii
Clade: Percomorpha
(unranked): Ovalentaria
Order: Cichliformes
Family: Cichlidae
Bonaparte,1835
Subfamilies and Tribes

Alternate taxonomy:

Forgenera,seebelow.

For the homophonous moth genus, seeCyclidia.

Cichlids/ˈsɪklɪdz/[a] arefishfrom thefamilyCichlidaein the orderCichliformes.Traditionally Cichlids were classed in a suborder, theLabroidei,along with the wrasses (Labridae), in the orderPerciformes,[3]but molecular studies have contradicted this grouping.[4]On the basis of fossil evidence, it first appeared inArgentinaduring theEarly Eoceneepoch, about 48.6 million years ago;[1]however,molecular clockestimates have placed the family's origin as far back as 67 million years ago, during thelate Cretaceousperiod.[2]The closest living relative of cichlids is probably theconvict blenny,and both families are classified in the 5th edition ofFishes of the Worldas the two families in the Cichliformes, part of the subseriesOvalentaria.[5]This family is large, diverse, and widely dispersed. At least 1,650specieshave beenscientifically described,[6]making it one of the largestvertebratefamilies. New species are discovered annually, and many species remainundescribed.The actual number of species is therefore unknown, with estimates varying between 2,000 and 3,000.[7]

Many cichlids, particularlytilapia,are important food fishes, while others, such as theCichlaspecies, are valued game fish. The family also includes many popularfreshwater aquariumfishkeptby hobbyists, including theangelfish,oscars,anddiscus.[8][9]Cichlids have the largest number ofendangered speciesamongvertebratefamilies, most in thehaplochrominegroup.[10]Cichlids are particularly well known for havingevolved rapidlyinto manyclosely related but morphologically diverse specieswithin large lakes, particularly LakesTanganyika,Victoria,Malawi,andEdward.[11][12]Their diversity in the African Great Lakes is important for the study ofspeciationin evolution.[13]Many cichlidsintroducedinto waters outside of their natural range have become nuisances.[14]

All cichlids practice some form of parental care for their eggs and fry, usually in the form of guarding the eggs and fry ormouthbrooding.

Anatomy and appearance

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Relationships within the Labrodei[3]

Cichlids span a wide range of body sizes, from species as small as 2.5 cm (1 in) in length (e.g., femaleNeolamprologus multifasciatus) to much larger species approaching 1 m (3 ft) in length (BoulengerochromisandCichla). As a group, cichlids exhibit a similar diversity of body shapes, ranging from strongly laterally compressed species (such asAltolamprologus,Pterophyllum,andSymphysodon) to species that are cylindrical and highly elongated (such asJulidochromis,Teleogramma,Teleocichla,Crenicichla,andGobiocichla).[8]Generally, however, cichlids tend to be of medium size, ovate in shape, and slightly laterally compressed, and generally similar to the North Americansunfishesin morphology, behavior, and ecology.[15]

Cichlids share a single key trait - the fusion of the lowerpharyngeal bonesinto 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 labor between the"true jaws"(mandibles) and the "pharyngeal jaws".Cichlids are efficient and often highly specialized feeders that capture and process a very wide variety of food items. This is assumed to be one reason why they are so diverse.[8]

The features that distinguish them from the other families in the Labroidei include:[16]

  • A single nostril on each side of the forehead, instead of two
  • No bony shelf below the orbit of the eye
  • Division of thelateral lineorgan 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 generaTeleogrammaandGobiocichla)
  • A distinctively shapedotolith
  • Thesmall intestine's left-side exit from thestomachinstead of its right side as in other Labroidei

Taxonomy

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Kullander(1998) recognizes eightsubfamiliesof cichlids: theAstronotinae,Cichlasomatinae,Cichlinae,Etroplinae,Geophaginae,Heterochromidinae,Pseudocrenilabrinae,andRetroculinae.[17]A ninth subfamily, thePtychochrominae,was later recognized by Sparks and Smith.[18]Cichlid taxonomy is still debated, and classification ofgeneracannot yet be definitively given. A comprehensive system of assigningspeciestomonophyleticgenera is still lacking, and there is not complete agreement on what genera should be recognized in this family.[19]

As an example of the classification problems, Kullander[20]placed the African genusHeterochromisphylogeneticallywithinNeotropicalcichlids, although later papers[citation needed]concluded otherwise. Other problems center upon the identity of the putative common ancestor for the Lake Victoria superflock (many closely related species sharing a single habitat), and the ancestral lineages of Lake Tanganyikan cichlids.[citation needed]

A 19th century watercolor painting of a pale flag cichlid.
A 19th century watercolor painting of a pale flag cichlid by Jacques Burkhardt.

Phylogeny derived from morphological characters shows differences at the genus level with phylogeny based ongenetic loci.[21]A consensus remains that the Cichlidae as a family are monophyletic.[22]

In cichlid taxonomy,dentitionwas formerly used as a classifying characteristic, but this was complicated because in many cichlids, tooth shapes change with age, due to wear, and cannot be relied upon.Genome sequencingand other technologies transformed cichlid taxonomy.

Alternatively, all cichlid species native to thenew world,can be classified under the subfamily Cichlinae, while Etroplinae can classify all cichlid species native to theold world.

Distribution and habitat

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Pelmatolapia mariae,caught on a hook and line, in Australia: Originally from Africa, the species established feral populations in Australia.[23]

Cichlids are one of the largest vertebrate families in the world. They are most diverse in Africa and South America. Africa alone is host to at least an estimated 1,600 species.[19]Central AmericaandMexicohave about 120 species, as far north as theRio GrandeinSouth Texas.Madagascarhas its own distinctive species (Katria,Oxylapia,Paratilapia,Paretroplus,Ptychochromis,andPtychochromoides), only distantly related to those on the African mainland.[16][24]Native cichlids are largely absent in Asia, except for 9 species inIsrael,Lebanon,andSyria(Astatotilapia flaviijosephi,Oreochromis aureus,O. niloticus,Sarotherodon galilaeus,Coptodon zillii,andTristramellaspp.), two inIran(Iranocichla), and three inIndiaandSri Lanka(EtroplusandPseudetroplus).[19]If disregardingTrinidad and Tobago(where the few native cichlids are members of genera that are widespread in the South American mainland), the three species from the genusNandopsisare the only cichlids from theAntillesin theCaribbean,specificallyCubaandHispaniola.Europe, Australia, Antarctica, and North America north of the Rio Grande drainage have no native cichlids, although inFlorida,Hawaii,Japan, northern Australia, and elsewhere, feral populations of cichlids have become established asexotics.[23][25][26][27][28][29][30]

Although most cichlids are found at relatively shallow depths, several exceptions do exist. The deepest known occurrences areTrematocaraat more than 300 m (1,000 ft) below the surface inLake Tanganyika.[31]Others found in relatively deep waters include species such asAlticorpus macrocleithrumandPallidochromis tokoloshdown to 150 m (500 ft) below the surface in Lake Malawi,[32][33]and the whitish (nonpigmented) and blindLamprologus lethops,which is believed to live as deep as 160 m (520 ft) below the surface in theCongo River.[34]

Cichlids are less commonly found inbrackishandsaltwaterhabitats, though many species tolerate brackish water for extended periods;Mayaheros urophthalmus,for example, is equally at home in freshwatermarshesandmangroveswamps, and lives and breeds in saltwater environments such as the mangrove belts aroundbarrier islands.[8]Several species ofTilapia,Sarotherodon,andOreochromisareeuryhalineand can disperse along brackish coastlines between rivers.[19]Only a few cichlids, however, inhabit primarily brackish or salt water, most notablyEtroplus maculatus,Etroplus suratensis,andSarotherodon melanotheron.[35]The perhaps most extreme habitats for cichlids are the warmhypersaline lakeswhere the members of the generaAlcolapiaandDanakiliaare found.Lake AbaededinEritreaencompasses the entire distribution ofD. dinicolai,and its temperature ranges from 29 to 45 °C (84 to 113 °F).[36]

With the exception of the species from Cuba, Hispaniola, and Madagascar, cichlids have not reached any oceanic island and have a predominantlyGondwanandistribution, showing the precise sister relationships predicted byvicariance:Africa-South America and India-Madagascar.[37]The dispersal hypothesis, in contrast, requires cichlids to have negotiated thousands of kilometers of open ocean between India and Madagascar without colonizing any other island, or for that matter, crossing theMozambique Channelto Africa. Although the vast majority of Malagasy cichlids are entirely restricted to fresh water,Ptychochromis grandidieriandParetroplus polyactisare commonly found in coastal brackish water and are apparently salt tolerant,[38][39]as is also the case forEtroplus maculatusandE. suratensisfrom India and Sri Lanka.[40][41]

Ecology

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Feeding

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Within the cichlid family, carnivores, herbivores, omnivores, planktivores, and detritivores are known, meaning the Cichlidae encompass essentially the full range of food consumption possible in the animal kingdom. Various species have morphological adaptations for specific food sources,[42]but most cichlids consume a wider variety of foods based on availability. Carnivorous cichlids can be further divided into piscivorous and molluscivorous, since the morphology and hunting behavior differ greatly between the two categories. Piscivorous cichlids eat other fish, fry, larvae, and eggs. Some species eat the offspring of mouthbrooders by head-ramming, wherein the hunter shoves its head into the mouth of a female to expel her young and eat them.[43]Molluscivorous cichlids have several hunting strategies amongst the varieties within the group. Lake Malawi cichlids consume substrate and filter it out through their gill rakers to eat the mollusks that were in the substrate. Gill rakers are finger-like structures that line the gills of some fish to catch any food that might escape through their gills.[44]

The bumblebee cichlid,Pseudotropheus crabro,is specialised in feeding on parasites from the catfishBagrus meridionalis.[45]

Many cichlids are primarilyherbivores,feeding onalgae(e.g.Petrochromis) andplants(e.g.Etroplus suratensis). Small animals, particularlyinvertebrates,are only a minor part of their diets.

Other cichlids aredetritivoresand eat organic material, calledAufwuchs(offal); among these species are thetilapiinesof the generaOreochromis,Sarotherodon,andTilapia.

Other cichlids arepredatoryand eat little or no plant matter. These include generalists that catch a variety of small animals, including other fishes andinsectlarvae(e.g.Pterophyllum), as well as variety of specialists.Trematocranusis a specializedsnail-eater, whilePungu maclarenifeeds onsponges.A number of cichlids feed on other fish, either entirely or in part.Crenicichlaspecies are stealth predators that lunge from concealment at passing small fish, whileRhamphochromisspecies are open-water pursuit predators that chase down their prey.[46]Paedophagouscichlids such as theCaprichromisspecies eat other species' eggs or young, in some cases ramming the heads of mouthbrooding species to force them to disgorge their young.[47][48][49][50]Among the more unusual feeding strategies are those ofCorematodus,Docimodus evelynae,Plecodus,Perissodus,andGenyochromisspp., which feed on scales and fins of other fishes, a behavior known aslepidophagy,[51][52][53]along with the death-mimicking behaviour ofNimbochromisandParachromisspecies, which lay motionless, luring small fish to their side prior to ambush.[54][55]

This variety of feeding styles has helped cichlids to inhabit similarly varied habitats. Its pharyngeal teeth (in the throat) afford cichlids so many "niche" feeding strategies, because the jaws pick and hold food, while the pharyngeal teeth crush the prey.

Behavior

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Aggression

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Aggressive behavior in cichlids is ritualized and consists of multiple displays used to seek confrontation while being involved in evaluation of competitors,[56]coinciding with temporal proximity to mating. Displays ofritualized aggressionin cichlids include a remarkably rapid change in coloration, during which a successfully dominant[56]territorial male assumes a more vivid and brighter coloration, while a subordinate or "nonterritorial" male assumes a dull-pale coloration.[57]In addition to color displays, cichlids employ theirlateral linesto sense movements of water around their opponents to evaluate the competing male for physical traits/fitness.[58]Male cichlids are very territorial due to the pressure of reproduction, and establish their territory andsocial statusby physically driving out[59]challenging males (novel intruders)[60]through lateral displays (parallel orientation, uncovering gills),[61]biting, or mouth fights (head-on collisions of open mouths, measuring jaw sizes, and biting each other's jaws). The cichlid socialdichotomyis composed of a single dominant with multiple subordinates, where the physical aggression of males becomes a contest for resources[59](mates, territory, food). Female cichlids prefer to mate with a successfullyalphamale with vivid coloration, whose territory has food readily available.

Mating

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Cichlids mate eithermonogamouslyorpolygamously.[8]The mating system of a given cichlid species is not consistently associated with its brooding system. For example, although most monogamous cichlids are not mouthbrooders,Chromidotilapia,Gymnogeophagus,Spathodus,andTanganicodusall include – or consist entirely of – monogamous mouthbrooders. In contrast, numerous open- or cave-spawning cichlids are polygamous; examples include manyApistogramma,Lamprologus,Nannacara,andPelvicachromisspecies.[8][62]

Most adult male cichlids, specifically in the cichlid tribe Haplochromini, exhibit a unique pattern of oval-shaped color dots on their anal fins. These phenomena, known as egg spots, aid in the mouthbrooding mechanisms of cichlids. The egg spots consist of carotenoid-based pigment cells, which indicate a high cost to the organism, when considering that fish are not able to synthesize their own carotenoids.[63]

The mimicry of egg spots is used by males for the fertilization process. Mouthbrooding females lay eggs and immediately snatch them up with their mouths. Over millions of years, male cichlids have evolved egg spots to initiate the fertilization process more efficiently.[64]When the females are snatching up the eggs into their mouth, the males gyrate their anal fins, which illuminates the egg spots on his tail. Afterwards, the female, believing these are her eggs, places her mouth to the anal fin (specifically the genital papilla) of the male, which is when he discharges sperm into her mouth and fertilizes the eggs.[63]

The genuine color of egg spots is a yellow, red, or orange inner circle with a colorless ring surrounding the shape. Through phylogenetic analysis, using the mitochondrialND2gene, the true egg spots are thought to have evolved in the common ancestor of theAstatoreochromislineage and the modernHaplochrominisspecies. This ancestor was most likely riverine in origin, based on the most parsimonious representation of habitat type in the cichlid family.[65]The presence of egg spots in a turbid riverine environment would seem particularly beneficial and necessary for intraspecies communication.[65]

Two pigmentation genes are found to be associated with egg-spot patterning and color arrangement. These arefhl2-aandfhl2-b,which are paralogs.[64]These genes aid in pattern formation and cell-fate determination in early embryonic development. The highest expression of these genes was temporally correlated with egg-spot formation. A short, interspersed, repetitive element was also seen to be associated with egg spots. Specifically, it was evident upstream of the transcriptional start site offhl2in onlyHaplochrominisspecies with egg spots[64]

Self-fertilization

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The cichlidBenitochromisnigrodorsalisfrom Western Africa ordinarily undergoes biparental reproduction, but is also able to undergofacultative(optional) selfing (self-fertilization).[66]Facultative selfing may be an adaptive option when a mating partner is unavailable.[66]

Brood care

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Further information:List of fish species that protect their young

Pit spawning in cichlids

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Pit spawning, also referred to as substrate breeding, is a behavior in cichlid fish in which a fish builds a pit in the sand or ground, where a pair court and consequently spawn.[67]Many different factors go into this behavior of pit spawning, including female choice of the male and pit size, as well as the male defense of the pits once they are dug in the sand.[68]

Cichlids are often divided into two main groups: mouthbrooders and substrate brooders. Different parenting investment levels and behaviors are associated with each type of reproduction.[69]As pit spawning is a reproductive behavior, many different physiological changes occur in the cichlid while this process is occurring that interfere with social interaction.[70]Different kinds of species that pit spawn, and many different morphological changes occur because of this behavioral experience.[67]

Pit spawning is an evolved behavior across the cichlid group. Phylogenetic evidence from cichlids in Lake Tanganyika could be helpful in uncovering the evolution of their reproductive behaviors.[71]Several important behaviors are associated with pit spawning, including parental care, food provisioning,[72]and brood guarding.[73]

Mouth brooding vs. pit spawning

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One of the differences studied in African cichlids is reproductive behavior. Some species pit spawn and some are known as mouth brooders. Mouthbrooding is a reproductive technique where the fish scoop up eggs and fry for protection.[69]While this behavior differs from species to species in the details, the general basis of the behavior is the same. Mouthbrooding also affects how they choose their mates and breeding grounds. In a 1995 study, Nelson found that in pit-spawning females choose males for mating based on the size of the pit that they dig, as well as some of the physical characteristics seen in the males.[68]Pit spawning also differs from mouth brooding in the size and postnatal care exhibited. Eggs that have been hatched from pit-spawning cichlids are usually smaller than those of mouthbrooders. Pit-spawners' eggs are usually around 2 mm, while mouthbrooders are typically around 7 mm. While different behaviors take place postnatally between mouthbrooders and pit spawners, some similarities exist. Females in both mouthbrooders and pit-spawning cichlids take care of their young after they are hatched. In some cases, both parents exhibit care, but the female always cares for the eggs and newly hatched fry.[74]

Pit spawning process

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Many species of cichlids use pit spawning, but one of the less commonly studied species that exhibits this behavior is the NeotropicalCichlasoma dimerus.This fish is a substrate breeder that displays biparental care after the fry have hatched from their eggs. One study[67]examined reproductive and social behaviors of this species to see how they accomplished their pit spawning, including different physiological factors such as hormone levels, color changes, and plasma cortisol levels. The entire spawning process could take about 90 minutes and 400~800 eggs could be laid. The female deposits about 10 eggs at a time, attaching them to the spawning surface, which may be a pit constructed on the substrate or another surface. The number of eggs laid was correlated to the space available on the substrate. Once the eggs were attached, the male swam over the eggs and fertilized them. The parents would then dig pits in the sand, 10–20 cm wide and 5–10 cm deep, where larvae were transferred after hatching. Larvae began swimming 8 days after fertilization and parenting behaviors and some of the physiological factors measured changed.

Color changes

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In the same study, color changes were present before and after the pit spawning occurred. For example, after the larvae were transferred and the pits were beginning to be protected, their fins turned a dark grey color.[67]In another study, of the rainbow cichlid,Herotilapia multispinosa,[70]color changes occurred throughout the spawning process. Before spawning, the rainbow cichlid was an olive color with grey bands. Once spawning behaviors started, the body and fins of the fish became a more golden color. When the eggs were finished being laid, the pelvic fin all the way back to the caudal fin turned to a darker color and blackened in both the males and the females.[70]

Pit sizes

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Females prefer a bigger pit size when choosing where to lay eggs.[68]Differences are seen in the sizes of pits that created, as well as a change in the morphology of the pits.[75]Evolutionary differences between species of fish may cause them to either create pits or castles when spawning. The differences were changes in the way that each species fed, their macrohabitats, and the abilities of their sensory systems.[75]

Evolution

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Cichlids are renowned for their recent, rapid evolutionary radiation, both across the entire clade and within different communities across separate habitats.[69][71][75][76][77][78]Within their phylogeny, many parallel instances are seen of lineages evolving to the same trait and multiple cases of reversion to an ancestral trait.

The family Cichlidae arose between 80 and 100 million years ago within the order Perciformes (perch-like fishes).[76]Cichlidae can be split into a few groups based on their geographic location: Madagascar, Indian, African, and Neotropical (or South American). The most famous and diverse group, the African cichlids, can be further split either into Eastern and Western varieties, or into groups depending on which lake the species is from:Lake Malawi,Lake Victoria,orLake Tanganyika.[76][77]Of these subgroups, the Madagascar and Indian cichlids are the most basal and least diverse.[citation needed]

Of the African cichlids, the West African or Lake Tanganyika cichlids are the most basal.[71][76] Cichlids' common ancestor is believed to have been a spit-spawning species.[77]Both Madagascar and Indian cichlids retain this feature. However, of the African cichlids, all extant substrate brooding species originate solely from Lake Tanganyika.[69][77]The ancestor of the Lake Malawi and Lake Victoria cichlids were mouthbrooders. Similarly, only around 30% of South American cichlids are thought to retain the ancestral substrate-brooding trait. Mouthbrooding is thought to have evolved individually up to 14 times, and a return to substrate brooding as many as three separate times between both African and Neotropical species.[77]

Associated behaviors

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Cichlids have a great variety of behaviors associated with substrate brooding, including courtship and parental care alongside the brooding and nest-building behaviors needed for pit spawning. Cichlids' behavior typically revolves around establishing and defending territories when not courting, brooding, or raising young. Encounters between males and males or females and females are agonistic, while an encounter between a male and female leads to courtship.[79]Courtship in male cichlids follows the establishment of some form of territory, sometimes coupled with building a bower to attract mates.[68][75][79]After this, males may attempt to attract female cichlids to their territories by a variety of lekking display strategies or otherwise seek out females of their species.[68]However, cichlids, at the time of spawning, undergo a behavioral change such that they become less receptive to outside interactions.[79]This is often coupled with some physiological change in appearance.[67][70][79]

Brood care

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Cichlids can have maternal, paternal, or biparental care. Maternal care is most common among mouthbrooders, but cichlids' common ancestor is thought to exhibit paternal-only care.[77]Other individuals outside of the parents may also play a role in raising young; in the biparental daffodil cichlid (Neolamprologus pulcher), closely related satellite males, those males that surround other males' territories and attempt to mate with female cichlids in the area, help rear the primary males' offspring and their own.[80]

A common form of brood care involves food provisioning. For example, females of lyretail cichlids (Neolamprologus modabu) dig at sandy substrate more to push nutritional detritus and zooplankton into the surrounding water. Adult ofN. modabuperform this strategy to collect food for themselves, but dig more when offspring are present, likely to feed their fry.[73][81]This substrate-disruption strategy is rather common and can also be seen in convict cichlids (Cichlasoma nigrofasciatum).[72][81]Other cichlids have an ectothermal mucus that they grow and feed to their young, while still others chew and distribute caught food to offspring. These strategies, however, are less common in pit-spawning cichlids.[81]

A substrate brooding female managuense cichlid,Parachromis managuense,guards a clutch of eggs in the aquarium.

Cichlids have highly organized breeding activities.[19]All species show some form of parental care for botheggsandlarvae,often nurturing free-swimming young until they are weeks or months old. Communal parental care, where multiple monogamous pairs care for a mixed school of young have also been observed in multiple cichlid species, includingAmphilophus citrinellus,Etroplus suratensis,andTilapia rendalli.[82][83][84]Comparably, the fry ofNeolamprologus brichardi,a species that commonly lives in large groups, are protected not only by the adults, but also by olderjuvenilesfrom previous spawns.[85]Several cichlids, including discus (Symphysodonspp.), someAmphilophusspecies,Etroplus,andUaruspecies, feed their young with a skinsecretionfrom mucous glands.[8][86]

The speciesNeolamprologus pulcheruses a cooperative breeding system, in which one breeding pair has many helpers that are subordinate to the dominant breeders.

Parental care falls into one of four categories:[86]substrate or open brooders, secretive cave brooders (also known as guarding speleophils[87]), and at least two types ofmouthbrooders,ovophile mouthbrooders and larvophile mouthbrooders.[88]

Open brooding

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Open- or substrate-brooding cichlids lay their eggs in the open, on rocks, leaves, or logs. Examples of open-brooding cichlids includePterophyllumandSymphysodonspecies andAnomalochromis thomasi.Male and female parents usually engage in differing brooding roles. Most commonly, the male patrols the pair's territory and repels intruders, while the female fans water over the eggs, removing the infertile ones, and leading the fry while foraging. Both sexes are able to perform the full range of parenting behaviours.[88]

Cave brooding

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A femaleCyphotilapia frontosamouthbrooding fry, which can be seen looking out her mouth

Secretive cave-spawning cichlids lay their eggs in caves, crevices, holes, ordiscarded mollusc shells,frequently attaching the eggs to the roof of the chamber. Examples includePelvicachromisspp.,Archocentrusspp., andApistogrammaspp.[86]Free-swimming fry and parents communicate in captivity and in the wild. Frequently, this communication is based on body movements, such as shaking andpelvic finflicking. In addition, open- and cave-brooding parents assist in finding food resources for their fry. Multiple neotropical cichlid species perform leaf-turning and fin-digging behaviors.[88]

Ovophile mouthbrooding

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Ovophile mouthbrooders incubate their eggs in their mouths as soon as they are laid, and frequently mouthbrood free-swimming fry for several weeks. Examples include manyEast African Riftlakes (Lake Malawi,Lake Tanganyika,andLake Victoria) endemics, e.g.:Maylandia,Pseudotropheus,Tropheus,andAstatotilapia burtoni,along with someSouth Americancichlids such asGeophagus steindachneri.

Larvophile mouthbrooding

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Larvophile mouthbrooders lay eggs in the open or in a cave and take the hatched larvae into the mouth. Examples include some variants ofGeophagus altifrons,and someAequidens,Gymnogeophagus,andSatanoperca,as well asOreochromis mossambicusandOreochromis niloticus.[8][86]Mouthbrooders, whether of eggs or larvae, are predominantly females. Exceptions that also involve the males include eretmodine cichlids (generaSpathodus,Eretmodus,andTanganicodus), someSarotherodonspecies (such asSarotherodon melanotheron[89]),Chromidotilapia guentheri,and someAequidensspecies.[8][88][90]This method appears to have evolved independently in several groups of African cichlids.[19]

Speciation

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Cichlids of theAfrican rift lake systemevolved from an originalhybrid swarm.[91]

Cichlids provide scientists with a unique perspective of speciation, having become extremely diverse in the recent geological past, those of Lake Victoria actually within the last 10,000 to 15,000 years, a small fraction of the millions taken for Galápagos finch speciation in Darwin's textbook case.[92]Some of the contributing factors to their diversification are believed to be the various forms of prey processing displayed by cichlidpharyngeal jawapparatus. These different jaw apparatus allow for a broad range of feeding strategies, including algae scraping, snail crushing, planktivory, piscivory, and insectivory.[93]Some cichlids can also showphenotypic plasticityin their pharyngeal jaws, which can also help lead to speciation. In response to different diets or food scarcity, members of the same species can display different jaw morphologies that are better suited to different feeding strategies. As species members begin to concentrate around different food sources and continue their lifecycle, they most likely spawn with like individuals. This can reinforce the jaw morphology and given enough time, create new species.[94]Such a process can happen throughallopatric speciation,whereby species diverge according to different selection pressures in different geographical areas, or throughsympatric speciation,by which new species evolve from a common ancestor while remaining in the same area. InLake ApoyoinNicaragua,Amphilophus zaliosusand its sister speciesAmphilophus citrinellusdisplay many of the criteria needed for sympatric speciation.[95]In theAfrican rift lake system,cichlid species in numerous distinct lakes evolved from a sharedhybrid swarm.[91]

Population status

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In 2010, theInternational Union for Conservation of Natureclassified 184 species asvulnerable,52 asendangered,and 106 ascritically endangered.[96] At present, the IUCN only listsYssichromissp. nov.argensasextinct in the wild,and six species are listed as entirely extinct, but many more possibly belong in these categories (for example,Haplochromis aelocephalus,H. apogonoides,H. dentex,H. dichrourus,and numerous other members of the genusHaplochromishave not been seen since the 1980s, but are maintained as critically endangered on the small chance that tiny –but currently unknown– populations survive).[96]

Lake Victoria

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Main article:Lake Victoria § Cichlid fish
Haplochromisthereuterionhas declined, but still survives in low numbers.[97]

Because of the introducedNile perch(Lates niloticus),Nile tilapia(Oreochromis niloticus), andwater hyacinth,deforestationthat led to watersiltation,andoverfishing,many Lake Victoria cichlid species have become extinct or been drastically reduced. By around 1980, lake fisheries yielded only 1% cichlids, a drastic decline from 80% in earlier years.[98]

By far the largest Lake Victoria group is the haplochromine cichlids, with more than 500 species, but at least 200 of these (about 40%) have become extinct,[99][100][101]and many others are seriously threatened.[102]Initially it was feared that the percentage of extinct species was even higher,[103]but some species have been rediscovered after the Nile perch started to decline in the 1990s.[100][104]Some species have survived in nearby small satellite lakes,[104]or in refugia among rocks orpapyrus sedges(protecting them from the Nile perch),[105]or have adapted to the human-induced changes in the lake itself.[100][101]The species were often specialists and these were not affected to the same extent. For example, thepiscivoroushaplochromines were particularly hard hit with a high number of extinctions,[106]while thezooplanktivoroushaplochromines reached densities in 2001 that were similar to before the drastic decline, although consisting of fewer species and with some changes in their ecology.[100]

Food and game fish

[edit]

Although cichlids are mostly small- to medium-sized, many are notable as food and game fishes. With few thick rib bones and tasty flesh,artisan fishingis not uncommon in Central America and South America, as well as areas surrounding the Africanrift lakes.[98]

Tilapia

[edit]

The most important food cichlids, however, are thetilapiinesof North Africa. Fast growing, tolerant of stocking density, and adaptable, tilapiine species have been introduced and farmed extensively in many parts of Asia and are increasingly commonaquaculturetargets elsewhere.

Farmed tilapia production is about 1,500,000 tonnes (1,700,000 short tons) annually, with an estimated value of US$1.8 billion,[107]about equal to that ofsalmonandtrout.

Unlike those carnivorous fish, tilapia can feed onalgaeor any plant-based food. This reduces the cost of tilapia farming, reduces fishing pressure on prey species, avoids concentrating toxins that accumulate at higher levels of thefood chain,and makes tilapia the preferred "aquatic chickens" of the trade.[98]

Game fish

[edit]

Many large cichlids are popular game fish. Thepeacock bass(Cichlaspecies) of South America is one of the most popularsportfish.It was introduced in many waters around the world.[where?]InFlorida,this fish generates millions of hours of fishing and sportfishing revenue of more than US$8 million a year.[108]Other cichlids preferred by anglers include theoscar,Mayan cichlid(Cichlasoma urophthalmus), andjaguar cichlid(Parachromis managuensis).[108]

Aquarium fish

[edit]
Further information:List of cichlid fish of South America
The discus,Symphysodonspp., has been popular among aquarium enthusiasts.

Since 1945, cichlids have become increasingly popular as aquarium fish.[8][86][88][109][110][111][112]

The most common species in hobbyist aquaria isPterophyllum scalarefrom theAmazon Riverbasin in tropical South America, known in the trade as the "angelfish".Other popular or readily available species include theoscar(Astronotus ocellatus),convict cichlid(Archocentrus nigrofasciatus) anddiscus fish(Symphysodon).[8]

Hybrids and selective breeding

[edit]
The "red Texas cichlid" is not aTexas cichlid(Herichthys cyanoguttatus) but a cross-genus hybrid ofHerichthysandAmphilophusparents.

Some cichlids readilyhybridizewith related species, both in the wild and under artificial conditions.[113]Other groups of fishes, such as Europeancyprinids,also hybridize.[114]Unusually, cichlid hybrids have been put to extensive commercial use, in particular for aquaculture and aquaria.[9][115]The hybrid red strain oftilapia,for example, is often preferred in aquaculture for its rapid growth. Tilapia hybridization can produce all-male populations to control stock density or prevent reproduction in ponds.[9]

Aquarium hybrids

[edit]

The most common aquarium hybrid is perhaps theblood parrot cichlid,which is a cross of several species, especially from species in the genusAmphilophus.(There are many hypotheses, but the most likely is:Amphilophus labiatus×Vieja synspillus[citation needed]With a triangular-shaped mouth, an abnormalspine,and an occasionally missingcaudal fin(known as the "love heart"parrotcichlid), the fish is controversial among aquarists. Some have called blood parrot cichlids "the Frankenstein monster of the fish world".[116]Another notable hybrid, theflowerhorn cichlid,was very popular in some parts ofAsiafrom 2001 until late 2003, and is believed to bring good luck to its owner.[117]The popularity of the flowerhorn cichlid declined in 2004.[118]Owners released many specimens into the rivers and canals ofMalaysiaandSingapore,where they threaten endemic communities.[119]

Aleucisticlong-finned form of the oscar,A. ocellatus

Numerous cichlid species have beenselectively bredto develop ornamental aquarium strains. The most intensive programs have involved angelfish and discus, and manymutationsthat affect both coloration and fins are known.[8][120][121]Other cichlids have been bred foralbino,leucistic,andxanthisticpigmentmutations,includingoscars,convict cichlidandPelvicachromis pulcher.[8][86]Bothdominantandrecessivepigment mutations have been observed.[122]Inconvict cichlids,for example, a leucistic coloration is recessively inherited,[123]while inOreochromis niloticus niloticus,red coloration is caused by a dominant inherited mutation.[124]

This selective breeding may haveunintended consequences.For example, hybrid strains ofMikrogeophagus ramirezihave health and fertility problems.[125]Similarly, intentionalinbreedingcan cause physical abnormalities, such as the notchedphenotypeinangelfish.[126]

Genera

[edit]

The genus list is as perFishBase.Studies are continuing, however, on the members of this family, particularly the haplochromine cichlids of the African rift lakes.[16]

[edit]

Footnotes

[edit]
  1. ^ Cichlidis frequently mispronounced in the pet trade as if spelled "chicklid"/ˈtʃɪklɪd/,presumably from confusion with names likeChiclets,and with Italian words likecioppinoandciaothat start withci-and the sound/tʃ/.

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