98-558: The damba ( Paretroplus damii ) is a species of cichlid . The genus name Paretroplus is composed by the Greek Para (meaning similar to ) and Etroplus (a closely related genus of Indian cichlids). The Latin species name damii honors the naturalist Douwe Casparus van Dam (1827-1898). P. damii is the largest Paretroplus , reaching the total length of about 17–40 centimetres (6.7–15.7 in). No sexual dimorphism exists in size or in other characteristics. These fishes have
196-419: 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
294-533: A genus level, six species of Chrysichthys catfish are only found in the Tanganyika basin where they live both in shallow and relatively deep waters; in the latter habitat they are the primary predators and scavengers. A unique evolutionary radiation in the lake is the 15 species of Mastacembelus spiny eels, all but one endemic to its basin. Although other African Great Lakes have Synodontis catfish, endemic catfish genera and Mastacembelus spiny eels,
392-502: A greater role than the rivers. At least 90% of the water influx is from rain falling on the lake's surface and at least 90% of the water loss is from direct evaporation. The major river flowing into the lake is the Ruzizi River , formed about 10,000 years ago, which enters the north of the lake from Lake Kivu . The Malagarasi River , which is Tanzania's second largest river, enters the east side of Lake Tanganyika. The Malagarasi
490-468: 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
588-401: 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
686-604: 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
784-480: 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
882-507: 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
980-586: A precious biological resource for the study of speciation and evolution . The cichlids of the African Great Lakes , including Tanganyika, represent the most diverse extent of adaptive radiation in vertebrates. Some of the endemic species do occur slightly into the upper Lukuga River (Lake Tanganyika's outflow), but any further distribution into the Congo River basin is stemmed purely by geography and chemistry—Tanganyika's lake environment
1078-442: 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
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#17327650702541176-416: A rounded shape, with a light pink / grey colouration in absence of vertical barring. This species can be found in several river basins in northwestern Madagascar . This includes populations in far northern Madagascar that some have speculated represented an undescribed species , but a comparison of specimens did not support this, instead maintaining them as part of P. damii . In contrast, populations in
1274-505: 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
1372-491: 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
1470-528: 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,
1568-499: 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
1666-504: 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
1764-514: 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
1862-469: 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
1960-691: Is an African Great Lake . It is the second-largest freshwater lake by volume and the second deepest , in both cases after Lake Baikal in Siberia . It is the world's longest freshwater lake. The lake is shared among four countries— Tanzania , the Democratic Republic of the Congo (the DRC), Burundi , and Zambia —with Tanzania (46%) and the DRC (40%) possessing the majority of the lake. It drains into
2058-717: 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
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#17327650702542156-544: 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
2254-424: Is deeper than any other cichlid known. Some of the deepwater genera (e.g., Bathybates , Gnathochromis , Hemibates and Xenochromis ) have been caught in places virtually devoid of oxygen, and how they are able to survive there is unclear. Tanganyika cichlids are generally benthic (found at or near the bottom) and/or coastally. No Tanganyika cichlids are truly pelagic and offshore, except for some of
2352-534: 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
2450-488: Is far more stable and consistent than the rapids and fast-flowing sections of the Congo River. Additionally, Lake Tanganyika's water is alkaline and of a higher pH (which the cichlids prefer), containing a higher calcium and mineral content than the Congo's acidic , sediment -rich waters, which collect much organic detritus from the surrounding rainforests . In areas of the Congo away from rapids or whitewater ,
2548-461: Is home to more than 80 species of non-cichlid fish and about 60% of these are endemic. The open waters of the pelagic zone are dominated by four non-cichlid species: Two species of "Tanganyika sardine" ( Limnothrissa miodon and Stolothrissa tanganicae ) form the largest biomass of fish in this zone, and they are important prey for the forktail lates ( Lates microlepis ) and sleek lates ( L. stappersii ). Two additional lates are found in
2646-478: Is older than Lake Tanganyika, and before the lake was formed, it probably was a headwater of the Lualaba River , the main Congo River headstream. The lake has a complex history of changing flow patterns, due to its high altitude, great depth, slow rate of refill, and mountainous location in a turbulently volcanic area that has undergone climate changes. Apparently, it has rarely in the past had an outflow to
2744-596: 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
2842-497: Is typically at depths around 100 m (330 ft) in the northern part of the lake and 240–250 m (790–820 ft) in the south. The oxygen-devoid deepest sections contain high levels of toxic hydrogen sulphide and are essentially lifeless, except for bacteria . Lake Tanganyika and its associated wetlands are home to Nile crocodiles (including famous giant Gustave ), Zambian hinged terrapins , serrated hinged terrapins , and pan hinged terrapins (last species not in
2940-482: Is unclear if they have a similar behavior. The facultative brood parasites often lay their eggs synchronously with mouthbroding cichlids. The cichlid pick up the eggs in their mouth as if they were their own. Once the catfish eggs hatch the young eat the cichlid eggs. Six catfish genera are entirely restricted to the lake basin: Bathybagrus , Dinotopterus , Lophiobagrus , Phyllonemus , Pseudotanganikallabes and Tanganikallabes . Although not endemic on
3038-648: The Congo River system and ultimately into the Atlantic Ocean . Lake Tanganyika is situated within the Albertine Rift , the western branch of the East African Rift , and is confined by the mountainous walls of the valley. It is the largest rift lake in Africa and the second-largest lake by volume in the world. It is the deepest lake in Africa and holds the greatest volume of fresh water on
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3136-676: The Mahanara River in northeastern Madagascar were formerly also included in P. damii , but these were described as a separate species, P. loisellei , in 2011, and populations in the Ankofia River basin in northwestern Madagascar (south of true P. damii ) appear to represent a closely related undescribed species. This Cichlidae -related article 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
3234-473: 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
3332-974: 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
3430-404: 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
3528-599: 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
3626-422: The mantle cavity of the unionid mussel Pleiodon spekei , making it one of only two known commensal species of freshwater shrimp (the other is the sponge-living Caridina spongicola from Lake Towuti , Indonesia). Among Rift Valley lakes , Lake Tanganyika far surpasses all others in terms of crustacean and freshwater snail richness (both in total number of species and number of endemics). For example,
3724-405: 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
3822-713: The piscivorous Bathybates . Two of these, B. fasciatus and B. leo , mainly feed on Tanganyika sardines . Tanganyika cichlids differ extensively in ecology , and include species that are herbivores , detritivores , planktivores , insectivores , molluscivores , scavengers , scale-eaters and piscivores. These dietary specializations, however, have been shown to be variable and subject to opportunistic changes. That is, many species of Tanganyikan cichlid with specialized diets showed opportunistic, episodic exploitation of Stolothrissa tanganicae and Limnothrissa miodon when prey concentrations were unusually high. The fishes' breeding behavior falls into two main groups:
3920-651: 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
4018-535: 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
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4116-542: 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
4214-647: The Niemba River, to maintain a flow. The lake may also have at times had different inflows and outflows; inward flows from a higher Lake Rukwa , access to Lake Malawi and an exit route to the Nile have all been proposed to have existed at some point in the lake's history. Lake Tanganyika is an ancient lake , one of only twenty more than a million years old. Its three basins, which in periods with much lower water levels were separate lakes, are of different ages. The central began to form 9–12 million years ago (Mya),
4312-561: 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
4410-450: The Tanganyika thalassoids, which are part of Prosobranchia , are endemic to the lake. Initially they were believed to be related to similar marine snails, but they are now known to be unrelated. Their appearance is now believed to be the result of the highly diverse habitats in Lake Tanganyika and evolutionary pressure from snail-eating fish and, in particular, Platythelphusa crabs. A total of 17 freshwater snail genera are endemic to
4508-552: The accumulated sediment and organic matter creates sections of " blackwater ", with a high concentration of tannins from dissolving wood and leaves, creating an environment in which cichlids simply do not thrive. Likewise, many tropical riverine species would likely suffer if exposed to the crisp, alkaline lake water. Although Lake Tanganyika has fewer cichlid species than Lakes Malawi or Victoria —which both have experienced relatively recent explosive species radiations (resulting in many closely related species)—, its cichlids are
4606-495: 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
4704-436: The area were fishing. Most of them included using a lantern as a lure for fish that are attracted to light. There were three basic forms. One called Lusenga which is a wide net used by one person from a canoe. The second one is using a lift net. This was done by dropping a net deep below the boat using two parallel canoes and then simultaneously pulling it up. The third is called Chiromila which consisted of three canoes. One canoe
4802-456: 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
4900-416: 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
4998-402: The continent, accounting for 16% of the world's available fresh water. It extends for 676 km (420 mi) in a general north–south direction and averages 50 km (31 mi) in width. The lake covers 32,900 km (12,700 sq mi), with a shoreline of 1,828 km (1,136 mi), a mean depth of 570 m (1,870 ft) and a maximum depth of 1,470 m (4,820 ft) (in
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#17327650702545096-892: 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
5194-527: 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,
5292-466: 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
5390-583: 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
5488-770: 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
5586-439: 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
5684-980: 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
5782-522: The fish populations, causing significant declines. In 2016, it was estimated that the total catch was up to 200,000 tonnes. It is thought that early Homo sapiens were making an impact on the region during the Stone Age . The time period of the Middle Stone Age to Late Stone Age is described as an age of advanced hunter-gatherers. There are many methods in which the native people of
5880-401: 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
5978-482: The genera Altolamprologus , Cyprichromis , Eretmodus , Julidochromis , Lamprologus , Neolamprologus , Tropheus and Xenotilapia , are popular aquarium fish due to their bright colors and patterns, and interesting behaviors. Recreating a Lake Tanganyika biotope to host those cichlids in a habitat similar to their natural environment is also popular in the aquarium hobby, with many species today being bred in captivity successfully. Lake Tanganyika
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#17327650702546076-453: 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
6174-594: 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
6272-608: The lake ( Haplochromini , Tilapiini and Tylochromini ). Others have proposed splitting the Tanganyika cichlids into as many as 12–16 tribes ( Bathybatini , Benthochromini , Boulengerochromini , Cyphotilapiini , Eretmodini , Greenwoodochromini , Perissodini and Trematocarini , in addition to the aforementioned tribes). Most Tanganyika cichlids live along the shoreline, down to a depth of 100 m (330 ft), but some deep-water species regularly descend to 200 m (660 ft). Trematocara species have, exceptionally, been found at more than 300 m (980 ft), which
6370-455: The lake itself, but in adjacent lagoons). Storm's water cobra , a threatened subspecies of banded water cobra that feeds mainly on fish, is only found in Lake Tanganyika, where it prefers rocky shores. Lake Tanganyika is home to at least 250 endemic species of cichlids , and undescribed species likely remain to be discovered. Almost all (roughly 98%) of the lake's cichlid species are found solely there, and nowhere else, thus making it
6468-579: The lake, and as a base from which to launch surprise attacks on Allied troops. It therefore became essential for the Allied forces to gain control of the lake themselves. Under the command of Lieutenant Commander Geoffrey Spicer-Simson the British Royal Navy achieved the monumental task of bringing two armed motor boats HMS Mimi and HMS Toutou from England to the lake by rail, road and river to Albertville (since renamed Kalemie in 1971) on
6566-936: The lake, such as Hirthia , Lavigeria , Paramelania , Reymondia , Spekia , Stanleya , Tanganyicia and Tiphobia . There are about 30 species of non-thalassoid snails in the lake, but only five of these are endemic, including Ferrissia tanganyicensis and Neothauma tanganyicense . The latter is the largest Tanganyika snail and its shell is often used by small shell-dwelling cichlids . Crustaceans are also highly diverse in Tanganyika with more than 200 species, of which more than half are endemic. They include 10 species of freshwater crabs (9 Platythelphusa and Potamonautes platynotus ; all endemic), at least 11 species of small atyid shrimp ( Atyella , Caridella and Limnocaridina ), an endemic palaemonid shrimp ( Macrobrachium moorei ), about 100 ostracods , including many endemics, and several copepods . Among these, Limnocaridina iridinae lives inside
6664-642: The lake, the Tanganyika lates ( L. angustifrons ) and bigeye lates ( L. mariae ), but both these are primarily benthic hunters, although they also may move into open waters. The four lates, all endemic to Tanganyika, have been overfished and larger individuals are rare today. Among the more unusual fish in the lake are the endemic, facultatively brood parasitic "cuckoo catfish", including at least Synodontis grandiops and S. multipunctatus . A number of others are very similar (e.g., S. lucipinnis and S. petricola ) and have often been confused; it
6762-412: The lake. Among the endemic bivalves are three monotypic genera: Grandidieria burtoni , Pseudospatha tanganyicensis and Brazzaea anceyi . Many of the snails are unusual for species living in freshwater in having noticeably thickened shells and/or distinct sculpture , features more commonly seen in marine snails. They are referred to as thalassoids, which can be translated to "marine-like". All
6860-626: The lake. He noted the name "Liemba" for its southern part, a word probably from the Fipa language . Tanganyika means "stars" in the Luvale language . The lake was the scene of Battle for Lake Tanganyika during World War I . With the aid of the Graf Goetzen , the Germans had complete control of the lake in the early stages of the war. The ship was used both to ferry cargo and personnel across
6958-493: 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
7154-505: The most morphologically and genetically diverse. This is linked to the maturity of Tanganyika, as it is far older than the other lakes. Tanganyika has the largest number of endemic cichlid genera of all African lakes. All Tanganyika cichlids are in the subfamily Pseudocrenilabrinae . Of the 10 tribes in this subfamily, half are largely or entirely restricted to the lake ( Cyprichromini , Ectodini , Lamprologini , Limnochromini and Tropheini ), while another three have species in
7252-460: The northern 7–8 Mya and the southern 2–4 Mya. The lake's water is alkaline with a pH around 9 at depths of 0–100 m (0–330 ft). Below this, it is around 8.7, gradually decreasing to 8.3–8.5 in the deepest parts of Tanganyika. A similar pattern can be seen in the electric conductivity , ranging from about 670 μS/cm in the upper part to 690 μS/cm in the deepest. Surface temperatures generally range from about 24 °C (75 °F) in
7350-503: The northern basin). It holds an estimated 18,750 km (4,500 cu mi). The catchment area of the lake is 231,000 km (89,000 sq mi). Two main rivers flow into the lake, as well as numerous smaller rivers and streams (whose lengths are limited by the steep mountains around the lake). The one major outflow is the Lukuga River , which empties into the Congo River drainage. Precipitation and evaporation play
7448-418: The only other Rift Valley lake with endemic freshwater crabs are Lake Kivu and Lake Victoria with two species each. The diversity of other invertebrate groups in Lake Tanganyika is often not well-known, but there are at least 20 described species of leeches (12 endemics), 9 sponges (7 endemic), 6 bryozoa (2 endemic), 11 flatworms (7 endemic), 20 nematodes (7 endemic), 28 annelids (17 endemic) and
7546-871: 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. Lake Tanganyika Lake Tanganyika ( / ˌ t æ ŋ ɡ ə n ˈ j iː k ə , - ɡ æ n -/ TANG -gən- YEE -kə, -gan- ; Kirundi : Ikiyaga ca Tanganyika )
7644-436: 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
7742-814: 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
7840-409: 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,
7938-648: 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
8036-421: 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
8134-404: 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
8232-663: The relatively high diversity is unique to Tanganyika, which likely is related to its old age. Among the non-endemic fish, some are widespread African species but several are only shared with the Malagarasi and Congo River basins, such as the Congo bichir ( Polypterus congicus ), goliath tigerfish ( Hydrocynus goliath ), Citharinus citharus , six-banded distichodus ( Distichodus sexfasciatus ) and mbu puffer ( Tetraodon mbu ). A total of 83 freshwater snail species (65 endemic) and 11 bivalve species (8 endemic) are known from
8330-470: The sea. It has been described as "practically endorheic " for this reason. The lake's connection to the sea is dependent on a high water level allowing water to overflow out of the lake through the Lukuga River into the Congo. When not overflowing, the lake's exit into the Lukuga River typically is blocked by sand bars and masses of weed, and instead this river depends on its own tributaries, especially
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
8526-413: The small hydrozoan jellyfish Limnocnida tanganyicae . Lake Tanganyika supports a major fishery, which, depending on source, provides 25–40% or c. 60% of the animal protein in the diet of the people living in the region. Lake Tanganyika fish can be found exported throughout East Africa. Major commercial fishing began in the mid-1950s and has, together with global warming, had a heavy impact on
8624-402: The south and is wind-driven, but to a lesser extent, up- and downwellings also occur elsewhere in the lake. As a consequence of the stratification, the deep sections contain " fossil water ". This also means it has no oxygen (it is anoxic ) in the deeper parts, essentially limiting fish and other aerobic organisms to the upper part. Some geographical variations are seen in this limit, but it
8722-579: The southern part of the lake in early August to 28–29 °C (82–84 °F) in the late rainy season in March—April. At depths greater than 400 m (1,300 ft), the temperature is very stable at 23.1–23.4 °C (73.6–74.1 °F). The water has gradually warmed since the 19th century and this has accelerated with global warming since the 1950s. The lake is stratified and seasonal mixing generally does not extend beyond depths of 150 m (490 ft). The mixing mainly occurs as upwellings in
8820-459: The substrate- or sand-spawners (often in caves or rock crevices) and the mouthbrooders . Among the endemic species are two of the world's smallest cichlids, Neolamprologus multifasciatus and N. similis (both shell dwellers ), measuring up to 4–5 cm (1.6–2.0 in), and one of the largest, the giant cichlid ( Boulengerochromis microlepis ) at up to 90 cm (3.0 ft). Many cichlids from Lake Tanganyika, such as species from
8918-645: 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,
9016-775: 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
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
9310-492: 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
9408-590: The western shore of Lake Tanganyika. The two boats waited until December 1915, and mounted a surprise attack on the Germans, with the capture of the gunboat Kingani . Another German vessel, the Hedwig , was sunk in February 1916, leaving the Götzen as the only German vessel remaining to control the lake. In order to avoid his prize ship falling into Allied hands, Zimmer scuttled the vessel on July 26, 1916. The vessel
9506-537: Was stationary with a lantern while another canoe holds one end of the net and the other circles the stationary one to meet up with the net. The first known Westerners to find the lake were the British explorers Richard Burton and John Speke , in 1858. They located it while searching for the source of the Nile River . Speke continued and found the actual source, Lake Victoria . Later David Livingstone passed by
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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