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98-531: Two species of cichlids are endemic to the river basin , but Ptychochromis onilahy is probably already extinct and the remaining range of Ptychochromoides betsileanus covers less than 10 square kilometres (3.9 sq mi). Sources of the Onilahy river are near Beadabo . It flows through Ankilimary , to Benenitra , Ehara , Bezaha and Antanimena . It is crossed by the RN 10 near Tameantsoa . The mouth of

196-506: A "species flock" of up to 1000 endemic species. Only seven cichlid species in Lake Malawi are not a part of the species flock: the Eastern happy ( Astatotilapia calliptera ), the sungwa ( Serranochromis robustus ), and five tilapia species (genera Oreochromis and Coptodon ). All of the other cichlid species in the lake are descendants of a single original colonist species, which itself

294-419: A behavioral change such that they become less receptive to outside interactions. This is often coupled with some physiological change in appearance. 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. Other individuals outside of the parents may also play a role in raising young; in

392-468: A group, cichlids exhibit a similar diversity of body shapes, ranging from strongly laterally compressed species (such as Altolamprologus , Pterophyllum , and Symphysodon ) to species that are cylindrical and highly elongated (such as Julidochromis , Teleogramma , Teleocichla , Crenicichla , and Gobiocichla ). Generally, however, cichlids tend to be of medium size, ovate in shape, and slightly laterally compressed, and generally similar to

490-401: A male and female leads to courtship. Courtship in male cichlids follows the establishment of some form of territory, sometimes coupled with building a bower to attract mates. 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. However, cichlids, at the time of spawning, undergo

588-604: A mating partner is unavailable. 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. 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. 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. As pit spawning

686-480: A minor part of their diets. Other cichlids are detritivores and eat organic material, called Aufwuchs (offal); among these species are the tilapiines of the genera Oreochromis , Sarotherodon , and Tilapia . Other cichlids are predatory and eat little or no 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

784-507: A mixed school of young have also been observed in multiple cichlid species, including Amphilophus citrinellus , Etroplus suratensis , and Tilapia rendalli . Comparably, the fry of Neolamprologus brichardi , a species that commonly lives in large groups, are protected not only by the adults, but also by older juveniles from previous spawns. Several cichlids, including discus ( Symphysodon spp.), some Amphilophus species, Etroplus , and Uaru species, feed their young with

882-509: A new environment. Any one of these ecological opportunities has the potential to result in an increase in population size and relaxed stabilizing (constraining) selection. As genetic diversity is positively correlated with population size the expanded population will have more genetic diversity compared to the ancestral population. With reduced stabilizing selection phenotypic diversity can also increase. In addition, intraspecific competition will increase, promoting divergent selection to use

980-442: A remarkably rapid change in coloration, during which a successfully dominant territorial male assumes a more vivid and brighter coloration, while a subordinate or "nonterritorial" male assumes a dull-pale coloration. In addition to color displays, cichlids employ their lateral lines to sense movements of water around their opponents to evaluate the competing male for physical traits/fitness. Male cichlids are very territorial due to

1078-505: A skin secretion from mucous glands. The species Neolamprologus pulcher uses 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: substrate or open brooders, secretive cave brooders (also known as guarding speleophils ), and at least two types of mouthbrooders , ovophile mouthbrooders and larvophile mouthbrooders. Open- or substrate-brooding cichlids lay their eggs in

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1176-491: 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

1274-517: A very wide variety of food items. This is assumed to be one reason why they are so diverse. The features that distinguish them from the other families in the Labroidei include: Kullander (1998) recognizes eight subfamilies of cichlids: the Astronotinae , Cichlasomatinae , Cichlinae , Etroplinae , Geophaginae , Heterochromidinae , Pseudocrenilabrinae , and Retroculinae . A ninth subfamily,

1372-418: A wider range of resources. This ecological release provides the potential for ecological speciation and thus adaptive radiation. Occupying a new environment might take place under the following conditions: A 2020 study found there to be no direct causal relationship between the proportionally most comparable mass radiations and extinctions in terms of "co-occurrence of species", substantially challenging

1470-499: 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. Molluscivorous cichlids have several hunting strategies amongst

1568-484: Is a process in which organisms diversify rapidly from an ancestral species into a multitude of new forms, particularly when a change in the environment makes new resources available, alters biotic interactions or opens new environmental niches . Starting with a single ancestor, this process results in the speciation and phenotypic adaptation of an array of species exhibiting different morphological and physiological traits. The prototypical example of adaptive radiation

1666-497: Is a reproductive behavior, many different physiological changes occur in the cichlid while this process is occurring that interfere with social interaction. Different kinds of species that pit spawn, and many different morphological changes occur because of this behavioral experience. Pit spawning is an evolved behavior across the cichlid group. Phylogenetic evidence from cichlids in Lake Tanganyika could be helpful in uncovering

1764-508: Is a specialized snail -eater, while Pungu maclareni feeds on sponges . A number of cichlids feed on other fish, either entirely or in part. Crenicichla species are stealth predators that lunge from concealment at passing small fish, while Rhamphochromis species are open-water pursuit predators that chase down their prey. Paedophagous cichlids such as the Caprichromis species eat other species' eggs or young, in some cases ramming

1862-518: Is a unique egg-brooding species, with 15 cm-long males amassing collections of shells and guarding them in the hopes of attracting females (about 6 cm in length) to lay eggs in these shells. These dominant males must defend their territories from three types of rival: (1) other dominant males looking to steal shells; (2) younger, "sneaker" males looking to fertilize eggs in a dominant male's territory; and (3) tiny, 2–4 cm "parasitic dwarf" males that also attempt to rush in and fertilize eggs in

1960-464: Is also the case for Etroplus maculatus and E. suratensis from India and Sri Lanka. 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, but most cichlids consume

2058-671: Is also the site of a separate major floral adaptive radiation event: the Hawaiian lobelioids . The Hawaiian lobelioids are significantly more speciose than the silverswords, perhaps because they have been present in Hawaii for so much longer: they descended from a single common ancestor who arrived in the archipelago up to 15 million years ago. Today the Hawaiian lobelioids form a clade of over 125 species, including succulents, trees, shrubs, epiphytes, etc. Many species have been lost to extinction and many of

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2156-717: Is based on body movements, such as shaking and pelvic fin flicking. 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. 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 many East African Rift lakes ( Lake Malawi , Lake Tanganyika , and Lake Victoria ) endemics, e.g.: Maylandia , Pseudotropheus , Tropheus , and Astatotilapia burtoni , along with some South American cichlids such as Geophagus steindachneri . Larvophile mouthbrooders lay eggs in

2254-479: Is believed to have been a spit-spawning species. Both Madagascar and Indian cichlids retain this feature. However, of the African cichlids, all extant substrate brooding species originate solely from Lake Tanganyika. 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

2352-405: Is believed to have formed 9–12 million years ago, putting a recent cap on the age of the lake's cichlid fauna. Many of Tanganyika's cichlids live very specialized lifestyles. The giant or emperor cichlid ( Boulengerochromis microlepis ) is a piscivore often ranked the largest of all cichlids (though it competes for this title with South America's Cichla temensis , the speckled peacock bass). It

2450-427: Is believed to have originated in Hawaii no more than 6 million years ago, making this one of Hawaii's youngest adaptive radiation events. This means that the silverswords evolved on Hawaii's modern high islands, and descended from a single common ancestor that arrived on Kauai from western North America. The closest modern relatives of the silverswords today are California tarweeds of the family Asteraceae . Hawaii

2548-534: Is equally at home in freshwater marshes and mangrove swamps, and lives and breeds in saltwater environments such as the mangrove belts around barrier islands . Several species of Tilapia , Sarotherodon , and Oreochromis are euryhaline and can disperse along brackish coastlines between rivers. Only a few cichlids, however, inhabit primarily brackish or salt water, most notably Etroplus maculatus , Etroplus suratensis , and Sarotherodon melanotheron . The perhaps most extreme habitats for cichlids are

2646-540: Is famously home to many piscivorous cichlid species, some of which feed by sucking the contents out of mouthbrooding females' mouths. Victoria's cichlids constitute a far younger radiation than even that of Lake Malawi, with estimates of the age of the flock ranging from 200,000 years to as little as 14,000. Hawaii has served as the site of a number of adaptive radiation events, owing to its isolation, recent origin, and large land area. The three most famous examples of these radiations are presented below, though insects like

2744-475: Is finch speciation on the Galapagos (" Darwin's finches "), but examples are known from around the world. Four features can be used to identify an adaptive radiation: Adaptive radiations are thought to be triggered by an ecological opportunity or a new adaptive zone. Sources of ecological opportunity can be the loss of antagonists ( competitors or predators ), the evolution of a key innovation, or dispersal to

2842-433: Is quite possibly the Hawaiian silverswords , named for alpine desert-dwelling Argyroxiphium species with long, silvery leaves that live for up to 20 years before growing a single flowering stalk and then dying. The Hawaiian silversword alliance consists of twenty-eight species of Hawaiian plants which, aside from the namesake silverswords, includes trees, shrubs, vines, cushion plants, and more. The silversword alliance

2940-465: Is scarce, the ground finches use their specialized beaks to eat the seeds that they are best suited to eat and thus avoid starvation. The other finches in the Galapagos are similarly uniquely adapted for their particular niche. The cactus finches ( Geospiza sp.) have somewhat longer beaks than the ground finches that serve the dual purpose of allowing them to feed on Opuntia cactus nectar and pollen while these plants are flowering, but on seeds during

3038-442: Is still an area of active research. One proposition is that the finches were able to have a non-adaptive, allopatric speciation event on separate islands in the archipelago, such that when they reconverged on some islands, they were able to maintain reproductive isolation . Once they occurred in sympatry, niche specialization was favored so that the different species competed less directly for resources. This second, sympatric event

Onilahy River - Misplaced Pages Continue

3136-499: Is thought that giant cichlids spawn only a single time, breeding in their third year and defending their young until they reach a large size, before dying of starvation some time thereafter. The three species of Altolamprologus are also piscivores, but with laterally compressed bodies and thick scales enabling them to chase prey into thin cracks in rocks without damaging their skin. Plecodus straeleni has evolved large, strangely curved teeth that are designed to scrape scales off of

3234-596: 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. 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

3332-546: Is used to probe the wood underneath for insects. Meanwhile, the ʻiʻiwi ( Drepanis coccinea ) has a very long curved beak for reaching nectar deep in Lobelia flowers. An entire clade of Hawaiian honeycreepers, the tribe Psittirostrini , is composed of thick-billed, mostly seed-eating birds, like the Laysan finch ( Telespiza cantans ). In at least some cases, similar morphologies and behaviors appear to have evolved convergently among

3430-482: The Greater Antilles ( Cuba , Hispaniola , Puerto Rico , and Jamaica ) have adaptively radiated in separate, convergent ways. On each of these islands, anoles have evolved with such a consistent set of morphological adaptations that each species can be assigned to one of six " ecomorphs ": trunk–ground, trunk–crown, grass–bush, crown–giant, twig, and trunk. Take for example crown–giants from each of these islands:

3528-466: The Ptychochrominae , was later recognized by Sparks and Smith. Cichlid taxonomy is still debated, and classification of genera cannot yet be definitively 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 classification problems, Kullander placed

3626-946: The Rio Grande in South Texas . Madagascar has its own distinctive species ( Katria , Oxylapia , Paratilapia , Paretroplus , Ptychochromis , and Ptychochromoides ), only distantly related to those on the African mainland. Native cichlids are largely absent in Asia, except for 9 species in Israel , Lebanon , and Syria ( Astatotilapia flaviijosephi , Oreochromis aureus , O. niloticus , Sarotherodon galilaeus , Coptodon zillii , and Tristramella spp.), two in Iran ( Iranocichla ), and three in India and Sri Lanka ( Etroplus and Pseudetroplus ). If disregarding Trinidad and Tobago (where

3724-404: The angelfish , oscars , and discus . Cichlids have the largest number of endangered species among vertebrate families, most in the haplochromine group. Cichlids are particularly well known for having evolved rapidly into many closely related but morphologically diverse species within large lakes, particularly Lakes Tanganyika , Victoria , Malawi , and Edward . Their diversity in

3822-583: 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 Argentina during the Early Eocene epoch, about 48.6 million years ago; however, molecular clock estimates have placed

3920-458: The island of Cocos south of Costa Rica ). Darwin's finches are not actually finches in the true sense, but are members of the tanager family Thraupidae , and are derived from a single ancestor that arrived in the Galapagos from mainland South America perhaps just 3 million years ago. Excluding the Cocos finch, each species of Darwin's finch is generally widely distributed in the Galapagos and fills

4018-454: The new world , can be classified under the subfamily Cichlinae, while Etroplinae can classify all cichlid species native to the old world . 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. Central America and Mexico have about 120 species, as far north as

Onilahy River - Misplaced Pages Continue

4116-642: The African Great Lakes is important for the study of speciation in evolution. Many cichlids introduced into waters outside of their natural range have become nuisances. All cichlids practice some form of parental care for their eggs and fry, usually in the form of guarding the eggs and fry or mouthbrooding . Cichlids span a wide range of body sizes, from species as small as 2.5 cm (1 in) in length (e.g., female Neolamprologus multifasciatus ) to much larger species approaching 1 m (3 ft) in length ( Boulengerochromis and Cichla ). As

4214-523: 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 , or Lake Tanganyika . Of these subgroups, the Madagascar and Indian cichlids are the most basal and least diverse. Of the African cichlids, the West African or Lake Tanganyika cichlids are the most basal. Cichlids' common ancestor

4312-529: The African genus Heterochromis phylogenetically within Neotropical cichlids, although later papers 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. Phylogeny derived from morphological characters shows differences at

4410-603: The Cuban Anolis luteogularis , Hispaniola's Anolis ricordii , Puerto Rico's Anolis cuvieri , and Jamaica's Anolis garmani (Cuba and Hispaniola are both home to more than one species of crown–giant). These anoles are all large, canopy-dwelling species with large heads and large lamellae (scales on the undersides of the fingers and toes that are important for traction in climbing), and yet none of these species are particularly closely related and appear to have evolved these similar traits independently. The same can be said of

4508-430: The Hawaiian drosophilid flies and Hyposmocoma moths have also undergone adaptive radiation. The Hawaiian honeycreepers form a large, highly morphologically diverse species group of birds that began radiating in the early days of the Hawaiian archipelago. While today only 17 species are known to persist in Hawaii (3 more may or may not be extinct), there were more than 50 species prior to Polynesian colonization of

4606-533: The Hawaiian honeycreepers; for example, the short, pointed beaks of Loxops and Oreomystis evolved separately despite once forming the justification for lumping the two genera together. The Hawaiian honeycreepers are believed to have descended from a single common ancestor some 15 to 20 million years ago, though estimates range as low as 3.5 million years. Adaptive radiation is not a strictly vertebrate phenomenon, and examples are also known from among plants. The most famous example of adaptive radiation in plants

4704-489: The Malawi species flock are mouth-brooders , meaning the female keeps her eggs in her mouth until they hatch; in almost all species, the eggs are also fertilized in the female's mouth, and in a few species, the females continue to guard their fry in their mouth after they hatch. Males of most species display predominantly blue coloration when mating. However, a number of particularly divergent species are known from Malawi, including

4802-554: The North American sunfishes in morphology, behavior, and ecology. Cichlids share 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 labor between the "true jaws" ( mandibles ) and the " pharyngeal jaws ". Cichlids are efficient and often highly specialized feeders that capture and process

4900-740: The Onilahy river is situated in the Indian Ocean at Saint Augustin, Madagascar , 35 km south of Toliara (Tuléar). Its main affluentes from its south are Sakamena river , Evasy River , Ianapera River , Isoanala river and the Ihazofotsy River . From the north these are Sakondry , Taheza , Sakamare and the Imatoto rivers. This article related to a river in Madagascar is a stub . You can help Misplaced Pages by expanding it . Cichlid Alternate taxonomy: For genera , see below . Cichlids / ˈ s ɪ k l ɪ d z / are fish from

4998-487: The anal fin (specifically the genital papilla) of the male, which is when he discharges sperm into her mouth and fertilizes the eggs. 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 mitochondrial ND2 gene, the true egg spots are thought to have evolved in the common ancestor of the Astatoreochromis lineage and

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5096-422: The archipelago (between 18 and 21 species have gone extinct since the discovery of the islands by westerners). The Hawaiian honeycreepers are known for their beaks, which are specialized to satisfy a wide range of dietary needs: for example, the beak of the ʻakiapōlāʻau ( Hemignathus wilsoni ) is characterized by a short, sharp lower mandible for scraping bark off of trees, and the much longer, curved upper mandible

5194-456: 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. 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

5292-416: 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. The mimicry of egg spots is used by males for

5390-548: The collective biomass of the Victoria cichlid species flock has decreased substantially and an unknown number of species have become extinct. However, the original range of morphological and behavioral diversity seen in the lake's cichlid fauna is still mostly present today, if endangered. These again include cichlids specialized for niches across the trophic spectrum, as in Tanganyika and Malawi, but again, there are standouts. Victoria

5488-563: The colubroid faunas in the rest of the world. These Madagascan examples are significantly older than most of the other examples presented here: Madagascar's fauna has been evolving in isolation since the island split from India some 88 million years ago, and the Mantellidae originated around 50 mya. Older examples are known: the K-Pg extinction event , which caused the disappearance of the dinosaurs and most other reptilian megafauna 65 million years ago,

5586-892: 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. 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

5684-420: The dominant male's territory. These parasitic dwarf males never grow to the size of dominant males, and the male offspring of dominant and parasitic dwarf males grow with 100% fidelity into the form of their fathers. A number of other highly specialized Tanganyika cichlids exist aside from these examples, including those adapted for life in open lake water up to 200m deep. The cichlids of Lake Malawi constitute

5782-527: The entire clade and within different communities across separate habitats. 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). 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,

5880-466: The evolution of their reproductive behaviors. Several important behaviors are associated with pit spawning, including parental care, food provisioning, and brood guarding. 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. While this behavior differs from species to species in

5978-676: The family Vangidae are marked by very distinct beak shapes to suit their ecological roles. Madagascan mantellid frogs have radiated into forms that mirror other tropical frog faunas, with the brightly colored mantellas ( Mantella ) having evolved convergently with the Neotropical poison dart frogs of Dendrobatidae , while the arboreal Boophis species are the Madagascan equivalent of tree frogs and glass frogs . The pseudoxyrhophiine snakes of Madagascar have evolved into fossorial, arboreal, terrestrial, and semi-aquatic forms that converge with

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6076-573: 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

6174-760: The female always cares for the eggs and newly hatched fry. Many species of cichlids use pit spawning, but one of the less commonly studied species that exhibits this behavior is the Neotropical Cichlasoma dimerus . This fish is a substrate breeder that displays biparental care after the fry have hatched from their eggs. One study 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

6272-439: 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. 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

6370-952: The few native cichlids are members of genera that are widespread in the South American mainland), the three species from the genus Nandopsis are the only cichlids from the Antilles in the Caribbean , specifically Cuba and Hispaniola . Europe, Australia, Antarctica, and North America north of the Rio Grande drainage have no native cichlids, although in Florida , Hawaii , Japan, northern Australia, and elsewhere, feral populations of cichlids have become established as exotics . Although most cichlids are found at relatively shallow depths, several exceptions do exist. The deepest known occurrences are Trematocara at more than 300 m (1,000 ft) below

6468-401: The full range of parenting behaviours. Secretive cave-spawning cichlids lay their eggs in caves, crevices, holes, or discarded mollusc shells , frequently attaching the eggs to the roof of the chamber. Examples include Pelvicachromis spp., Archocentrus spp., and Apistogramma spp. Free-swimming fry and parents communicate in captivity and in the wild. Frequently, this communication

6566-453: The genus level with phylogeny based on genetic loci . A consensus remains that the Cichlidae as a family are monophyletic. In cichlid taxonomy, dentition was 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 sequencing and other technologies transformed cichlid taxonomy. Alternatively, all cichlid species native to

6664-594: The heads of mouthbrooding species to force them to disgorge their young. Among the more unusual feeding strategies are those of Corematodus , Docimodus evelynae , Plecodus , Perissodus , and Genyochromis spp., which feed on scales and fins of other fishes, a behavior known as lepidophagy , along with the death-mimicking behaviour of Nimbochromis and Parachromis species, which lay motionless, luring small fish to their side prior to ambush. This variety of feeding styles has helped cichlids to inhabit similarly varied habitats. Its pharyngeal teeth (in

6762-459: The hypothesis of "creative mass extinctions". Darwin's finches on the Galapagos Islands are a model system for the study of adaptive radiation. Today represented by approximately 15 species, Darwin's finches are Galapagos endemics famously adapted for a specialized feeding behavior (although one species, the Cocos finch ( Pinaroloxias inornata ), is not found in the Galapagos but on

6860-529: The lakes (meaning that sympatric speciation was the most probable mechanism for initial specialization). Also, continual changes in the water level of the lakes during the Pleistocene (which often turned the largest lakes into several smaller ones) could have created the conditions for secondary allopatric speciation. Lake Tanganyika is the site from which nearly all the cichlid lineages of East Africa (including both riverine and lake species) originated. Thus,

6958-489: 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. Many cichlids, particularly tilapia , are important food fishes, while others, such as the Cichla species, are valued game fish. The family also includes many popular freshwater aquarium fish kept by hobbyists, including

7056-602: The modern Haplochrominis species. This ancestor was most likely riverine in origin, based on the most parsimonious representation of habitat type in the cichlid family. The presence of egg spots in a turbid riverine environment would seem particularly beneficial and necessary for intraspecies communication. Two pigmentation genes are found to be associated with egg-spot patterning and color arrangement. These are fhl2-a and fhl2-b , which are paralogs. These genes aid in pattern formation and cell-fate determination in early embryonic development. The highest expression of these genes

7154-763: The open or in a cave and take the hatched larvae into the mouth. Examples include some variants of Geophagus altifrons , and some Aequidens , Gymnogeophagus , and Satanoperca , as well as Oreochromis mossambicus and Oreochromis niloticus . Mouthbrooders, whether of eggs or larvae, are predominantly females. Exceptions that also involve the males include eretmodine cichlids (genera Spathodus , Eretmodus , and Tanganicodus ), some Sarotherodon species (such as Sarotherodon melanotheron ), Chromidotilapia guentheri , and some Aequidens species. This method appears to have evolved independently in several groups of African cichlids. Adaptive radiation In evolutionary biology , adaptive radiation

7252-436: The open, on rocks, leaves, or logs. Examples of open-brooding cichlids include Pterophyllum and Symphysodon species and Anomalochromis 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

7350-535: The other five ecomorphs across the Caribbean's four largest islands. Much like in the case of the cichlids of the three largest African Great Lakes, each of these islands is home to its own convergent Anolis adaptive radiation event. Presented above are the most well-documented examples of modern adaptive radiation, but other examples are known. Populations of three-spined sticklebacks have repeatedly diverged and evolved into distinct ecotypes. On Madagascar , birds of

7448-814: The physical aggression of males becomes a contest for resources (mates, territory, food). Female cichlids prefer to mate with a successfully alpha male with vivid coloration, whose territory has food readily available. Cichlids mate either monogamously or polygamously . 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 , and Tanganicodus all include – or consist entirely of – monogamous mouthbrooders. In contrast, numerous open- or cave-spawning cichlids are polygamous; examples include many Apistogramma , Lamprologus , Nannacara , and Pelvicachromis species. Most adult male cichlids, specifically in

7546-409: The physiological factors measured changed. 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. In another study, of the rainbow cichlid, Herotilapia multispinosa , color changes occurred throughout the spawning process. Before spawning,

7644-500: The piscivorous Nimbochromis livingtonii , which lies on its side in the substrate until small cichlids, perhaps drawn to its broken white patterning, come to inspect the predator - at which point they are swiftly eaten. Lake Victoria's cichlids are also a species flock, once composed of some 500 or more species. The deliberate introduction of the Nile Perch ( Lates niloticus ) in the 1950s proved disastrous for Victoria cichlids, and

7742-640: The precise sister relationships predicted by vicariance : Africa-South America and India-Madagascar. 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 the Mozambique Channel to Africa. Although the vast majority of Malagasy cichlids are entirely restricted to fresh water, Ptychochromis grandidieri and Paretroplus polyactis are commonly found in coastal brackish water and are apparently salt tolerant, as

7840-421: The pressure of reproduction, and establish their territory and social status by physically driving out challenging males (novel intruders) through lateral displays (parallel orientation, uncovering gills), biting, or mouth fights (head-on collisions of open mouths, measuring jaw sizes, and biting each other's jaws). The cichlid social dichotomy is composed of a single dominant with multiple subordinates, where

7938-404: 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. Females prefer a bigger pit size when choosing where to lay eggs. Differences are seen in

8036-404: The rest of the year. The warbler-finches ( Certhidea sp.) have short, pointed beaks for eating insects. The woodpecker finch ( Camarhynchus pallidus ) has a slender beak which it uses to pick at wood in search of insects; it also uses small sticks to reach insect prey inside the wood, making it one of the few animals that use tools . The mechanism by which the finches initially diversified

8134-482: The roles typically filled by many fish families, including those of predators, scavengers, and herbivores, with varying dentitions and head shapes to match their dietary habits. In each case, the radiation events are only a few million years old, making the high level of speciation particularly remarkable. Several factors could be responsible for this diversity: the availability of a multitude of niches probably favored specialization, as few other fish taxa are present in

8232-401: The same niche on each island. For the ground finches, this niche is a diet of seeds, and they have thick bills to facilitate the consumption of these hard materials. The ground finches are further specialized to eat seeds of a particular size: the large ground finch ( Geospiza magnirostris ) is the largest species of Darwin's finch and has the thickest beak for breaking open the toughest seeds,

8330-429: The sides of other fish, scales being its main source of food. Gnathochromis permaxillaris possesses a large mouth with a protruding upper lip, and feeds by opening this mouth downward onto the sandy lake bottom, sucking in small invertebrates. A number of Tanganyika's cichlids are shell-brooders, meaning that mating pairs lay and fertilize their eggs inside of empty shells on the lake bottom. Lamprologus callipterus

8428-408: The sizes of pits that created, as well as a change in the morphology of the pits. 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. Cichlids are renowned for their recent, rapid evolutionary radiation, both across

8526-484: The small ground finch ( Geospiza fuliginosa ) has a smaller beak for eating smaller seeds, and the medium ground finch ( Geospiza fortis ) has a beak of intermediate size for optimal consumption of intermediately sized seeds (relative to G. magnirostris and G. fuliginosa ). There is some overlap: for example, the most robust medium ground finches could have beaks larger than those of the smallest large ground finches. Because of this overlap, it can be difficult to tell

8624-415: The species apart by eye, though their songs differ. These three species often occur sympatrically , and during the rainy season in the Galapagos when food is plentiful, they specialize little and eat the same, easily accessible foods. It was not well-understood why their beaks were so adapted until Peter and Rosemary Grant studied their feeding behavior in the long dry season, and discovered that when food

8722-461: The species in the lake constitute a single adaptive radiation event but do not form a single monophyletic clade . Lake Tanganyika is also the least speciose of the three largest African Great Lakes, with only around 200 species of cichlid; however, these cichlids are more morphologically divergent and ecologically distinct than their counterparts in lakes Malawi and Victoria, an artifact of Lake Tanganyika's older cichlid fauna. Lake Tanganyika itself

8820-691: The surface in Lake Tanganyika . Others found in relatively deep waters include species such as Alticorpus macrocleithrum and Pallidochromis tokolosh down to 150 m (500 ft) below the surface in Lake Malawi, and the whitish (non pigmented ) and blind Lamprologus lethops , which is believed to live as deep as 160 m (520 ft) below the surface in the Congo River . Cichlids are less commonly found in brackish and saltwater habitats, though many species tolerate brackish water for extended periods; Mayaheros urophthalmus , for example,

8918-775: The surrounding water. Adult of N. modabu perform this strategy to collect food for themselves, but dig more when offspring are present, likely to feed their fry. This substrate-disruption strategy is rather common and can also be seen in convict cichlids ( Cichlasoma nigrofasciatum ). 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. Cichlids have highly organized breeding activities. All species show some form of parental care for both eggs and larvae , often nurturing free-swimming young until they are weeks or months old. Communal parental care, where multiple monogamous pairs care for

9016-544: The surviving species endangered. Anole lizards are distributed broadly in the New World, from the Southeastern US to South America. With over 400 species currently recognized, often placed in a single genus ( Anolis ), they constitute one of the largest radiation events among all lizards. Anole radiation on the mainland has largely been a process of speciation, and is not adaptive to any great degree, but anoles on each of

9114-406: The throat) afford cichlids so many "niche" feeding strategies, because the jaws pick and hold food, while the pharyngeal teeth crush the prey. Aggressive behavior in cichlids is ritualized and consists of multiple displays used to seek confrontation while being involved in evaluation of competitors, coinciding with temporal proximity to mating. Displays of ritualized aggression in cichlids include

9212-478: 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. Many cichlids are primarily herbivores , feeding on algae (e.g. Petrochromis ) and plants (e.g. Etroplus suratensis ). Small animals, particularly invertebrates , are only

9310-487: The warm hypersaline lakes where the members of the genera Alcolapia and Danakilia are found. Lake Abaeded in Eritrea encompasses the entire distribution of D. dinicolai , and its temperature ranges from 29 to 45 °C (84 to 113 °F). With the exception of the species from Cuba, Hispaniola, and Madagascar, cichlids have not reached any oceanic island and have a predominantly Gondwanan distribution, showing

9408-561: Was adaptive radiation. The haplochromine cichlid fishes in the Great Lakes of the East African Rift (particularly in Lake Tanganyika , Lake Malawi , and Lake Victoria ) form the most speciose modern example of adaptive radiation. These lakes are believed to be home to about 2,000 different species of cichlid, spanning a wide range of ecological roles and morphological characteristics. Cichlids in these lakes fill nearly all of

9506-416: Was descended from Tanganyikan ancestors. The common ancestor of Malawi's species flock is believed to have reached the lake 3.4 million years ago at the earliest, making Malawi cichlids' diversification into their present numbers particularly rapid. Malawi's cichlids span a similarly range of feeding behaviors to those of Tanganyika, but also show signs of a much more recent origin. For example, all members of

9604-520: 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 of fhl2 in only Haplochrominis species with egg spots The cichlid Benitochromis nigrodorsalis from Western Africa ordinarily undergoes biparental reproduction, but is also able to undergo facultative (optional) selfing ( self-fertilization ). Facultative selfing may be an adaptive option when

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