Template:Short description Template:Use dmy dates Template:Infobox body of water

File:View of Kagongo Ward.jpg
Lake Tanganyika eastern Shore in Kagongo Ward, Kigoma Region, Tanzania

Lake Tanganyika (Template:IPAc-en Template:Respell;<ref>Template:Cite LPD</ref> Template:Langx) is an African Great Lake.<ref>Template:Cite book</ref> It is the world's second-largest freshwater lake by volume and the second deepest, in both cases after Lake Baikal in Siberia.<ref name=zambiatour>{{#invoke:citation/CS1|citation |CitationClass=web }}</ref><ref>{{#invoke:citation/CS1|citation |CitationClass=web }}</ref> It is the world's longest freshwater lake.<ref name=zambiatour/> 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 via the Lukuga River into the Congo River system, which ultimately discharges at Banana, Democratic Republic of the Congo into the Atlantic Ocean.<ref name=asdf />

GeographyEdit

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 freshwater lake by volume in the world. It is the deepest lake in Africa and holds the greatest volume of fresh water on the continent, accounting for 16% of the world's available fresh water. It extends for Template:Cvt in a general north–south direction and averages Template:Cvt in width. The lake covers Template:Cvt, with a shoreline of Template:Cvt, a mean depth of Template:Cvt and a maximum depth of Template:Cvt (in the northern basin). It holds an estimated Template:Cvt of water.<ref>{{#invoke:citation/CS1|citation |CitationClass=web }}</ref><ref name=asdf />

The catchment area of the lake is Template:Cvt. 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 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.<ref name= Kullander2011>Template:Cite journal</ref>

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.<ref name="readersnatural">Template:Cite book</ref> The Malagarasi River, which is Tanzania's second largest river, enters the east side of Lake Tanganyika.<ref name="readersnatural" /> The Malagarasi 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.<ref name=Kullander2011/>

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 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.<ref name="readersnatural" /> 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 the Niemba River, to maintain a flow.<ref name=Kullander2011/>

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.<ref>Template:Cite book</ref>

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), the northern 7–8 Mya and the southern 2–4 Mya.<ref>Template:Cite journal</ref>

Water characteristicsEdit

File:Clear lake Kagongo Ward.jpg
Clear water lake of Lake Tanganyika in Kagongo Ward, Kigoma Region, Tanzania

The lake's water is alkaline with a pH around 9 at depths of Template:Cvt.<ref name=Wever2005>Template:Cite journal</ref> Below this, it is around 8.7, gradually decreasing to 8.3–8.5 in the deepest parts of Tanganyika.<ref name=Wever2005/> 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.<ref name=Wever2005/>

Surface temperatures generally range from about Template:Cvt in the southern part of the lake in early August to Template:Cvt in the late rainy season in March—April.<ref name=Edmond1993>Template:Cite journal</ref> At depths greater than Template:Cvt, the temperature is very stable at Template:Cvt.<ref name=Reilly2003>Template:Cite journal</ref> The water has gradually warmed since the 19th century and this has accelerated with global warming since the 1950s.<ref name=Jensen2016>{{#invoke:citation/CS1|citation |CitationClass=web }}</ref>

The lake is stratified and seasonal mixing generally does not extend beyond depths of Template:Cvt.<ref name=Edmond1993/> The mixing mainly occurs as upwellings in the south and is wind-driven, but to a lesser extent, up- and downwellings also occur elsewhere in the lake.<ref name=Lowe2003>Template:Cite journal</ref> As a consequence of the stratification, the deep sections contain "fossil water".<ref>Hutter; Yongqi; and Chubarenko (2011). Physics of Lakes, volume 1: Foundation of the Mathematical and Physical Background. P. 11. Template:ISBN.</ref> 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 is typically at depths around Template:Cvt in the northern part of the lake and Template:Cvt in the south.<ref name=Wright2006>Wright, J.J.; and L.M. Page (2006). Taxonomic revision of Lake Tanganyikan Synodontis (Siluriformes: Mochokidae). Florida Mus. Nat. Hist. Bull. 46(4): 99–154.</ref><ref name=McConnell1987>Lowe-McConnell, R.H. (1987). Ecological Studies in Tropical Fish Communities. Template:ISBN.</ref> The oxygen-devoid deepest sections contain high levels of toxic hydrogen sulfide and are essentially lifeless,<ref name=zambiatour/> except for bacteria.<ref name=Wever2005/><ref>Template:Cite journal</ref>

BiologyEdit

ReptilesEdit

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 the lake itself, but in adjacent lagoons).<ref name=Spawls2002>Spawls, Howell, Drewes, and Ashe (2002). A Field Guide to the Reptiles of East Africa. Academic Press, London. Template:ISBN.</ref> 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.<ref name=Spawls2002/><ref>Template:Cite journal</ref>

Cichlid fishesEdit

File:Murky waters.jpg
A biologist collecting samples in 2020 in the murky waters of Lake Tanganyika for a study on the diet adaptations of cichlid fishes to better understand their evolution and speciation.
File:Neolamprologus brichardi.jpg
One of the many Tanganyika cichlids is Neolamprologus brichardi. The complex behaviors of this species and its close relative N. pulcher have been studied in detail<ref>Template:Cite journal</ref><ref>Template:Cite journal</ref><ref>Template:Cite journal</ref>

Lake Tanganyika is home to at least 250 endemic species of cichlids,<ref name=LTBP>{{#invoke:citation/CS1|citation |CitationClass=web }}</ref> and undescribed species likely remain to be discovered.<ref name=MortiffTangDiv>{{#invoke:citation/CS1|citation |CitationClass=web }}</ref> Almost all (roughly 98%) of the lake's cichlid species are found solely there, and nowhere else, thus making it a precious biological resource for the study of speciation and evolution.<ref>Template:Cite journal</ref><ref>Template:Cite journal</ref> The cichlids of the African Great Lakes, including Tanganyika, represent the most diverse extent of adaptive radiation in vertebrates.<ref name=":0" /> 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 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, 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.<ref name="Kullander2011" /> 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)—,<ref name="Turner2001">Template:Cite journal</ref> its cichlids are the most morphologically and genetically diverse.<ref name=":0">Template:Cite journal</ref><ref>Template:Cite journal</ref> This is linked to the maturity of Tanganyika, as it is far older than the other lakes.<ref>Template:Cite journal</ref> Tanganyika has the largest number of endemic cichlid genera of all African lakes.<ref name=":0"/> 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 the lake (Haplochromini, Tilapiini and Tylochromini).<ref>Template:Cite journal</ref> 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).<ref name=":0"/>

Most Tanganyika cichlids live along the shoreline, down to a depth of Template:Cvt, but some deep-water species regularly descend to Template:Cvt.<ref name=Kirchberger2012>Template:Cite journal</ref> Trematocara species have, exceptionally, been found at more than Template:Cvt, which is deeper than any other cichlid known.<ref>Loiselle, Paul (1994). The Cichlid Aquarium, p. 304. Tetra Press, Germany. Template:ISBN.</ref> 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.<ref name=McConnell1987/> Tanganyika cichlids are generally benthic (found at or near the bottom) and/or coastally.<ref name=Lindqvist1999>Lindqvist, O.V.; H. Mölsä; K. Solonen; J. Sarvala, editors (1999). From Limnology to Fisheries: Lake Tanganyika and Other Large Lakes. pp. 213–214. Springer. Template:ISBN</ref> No Tanganyika cichlids are truly pelagic and offshore, except for some of the piscivorous Bathybates.<ref name=Kirchberger2012/> Two of these, B. fasciatus and B. leo, mainly feed on Tanganyika sardines.<ref name=Kirchberger2012/><ref name=McConnell1987/> Tanganyika cichlids differ extensively in ecology, and include species that are herbivores, detritivores, planktivores, insectivores, molluscivores, scavengers, scale-eaters and piscivores.<ref name=MortiffTangDiv/> 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.<ref>Template:Cite journal</ref> The fishes' breeding behavior falls into two main groups: the substrate- or sand-spawners (often in caves or rock crevices) and the mouthbrooders.<ref name=Schliewen1992>Schliewen, U. (1992). Aquarium Fish. Barron's Educational Series. Template:ISBN.</ref> Among the endemic species are two of the world's smallest cichlids, Neolamprologus multifasciatus and N. similis (both shell dwellers), measuring up to Template:Cvt,<ref>{{#invoke:Cite taxon|main|fishbase|genus=|species=|subspecies=}}</ref><ref>{{#invoke:Cite taxon|main|fishbase|genus=|species=|subspecies=}}</ref> and one of the largest, the giant cichlid (Boulengerochromis microlepis) at up to Template:Cvt.<ref name=MortiffTangDiv/><ref name=PracFishKeepLarge>{{#invoke:citation/CS1|citation |CitationClass=web }}</ref>

Many cichlids from Lake Tanganyika, such as species from 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.<ref name=Schliewen1992/> 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.<ref name=Schliewen1992/><ref>{{#invoke:citation/CS1|citation |CitationClass=web }}</ref>

Other fishEdit

Lake Tanganyika is home to more than 80 species of non-cichlid fish and about 60% of these are endemic.<ref name=Wright2006/><ref name=LTBP/><ref name=Brown2011>Template:Cite journal</ref><ref>Template:Cite journal</ref>

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).<ref name=Lindqvist1999/> Two additional lates are found in 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.<ref name=Lindqvist1999/> The four lates, all endemic to Tanganyika, have been overfished and larger individuals are rare today.<ref name=Lindqvist1999/>

Among the more unusual fish in the lake are the endemic, facultatively brood parasitic "cuckoo catfish", including at least Synodontis grandiops<ref>{{#invoke:citation/CS1|citation |CitationClass=web }}</ref> and S. multipunctatus.<ref name=Wright2006/><ref name=Schliewen1992/> A number of others are very similar (e.g., S. lucipinnis and S. petricola) and have often been confused; it is unclear if they have a similar behavior.<ref name=Wright2006/><ref>{{#invoke:citation/CS1|citation |CitationClass=web }}Template:Pb{{#invoke:citation/CS1|citation |CitationClass=web }}</ref> 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.<ref name=Wright2006/><ref name=Schliewen1992/> Six catfish genera are entirely restricted to the lake basin: Bathybagrus, Dinotopterus, Lophiobagrus, Phyllonemus, Pseudotanganikallabes and Tanganikallabes.<ref name=FishbaseList>{{#invoke:citation/CS1|citation |CitationClass=web }}</ref><ref>Template:Cite journal</ref> Although not endemic on 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;<ref name=FishbaseList/> in the latter habitat they are the primary predators and scavengers.<ref name=McConnell1987/> A unique evolutionary radiation in the lake is the 15 species of Mastacembelus spiny eels, all but one endemic to its basin.<ref name=Brown2011/><ref name=Brown2010>Template:Cite journal</ref> Although other African Great Lakes have Synodontis catfish, endemic catfish genera and Mastacembelus spiny eels, the relatively high diversity is unique to Tanganyika, which likely is related to its old age.<ref name=Brown2010/>

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).<ref name=FishbaseList/>

Molluscs and crustaceansEdit

File:Tiphobia horei 01.JPG
The shell of the endemic thalassoid freshwater snail Tiphobia horei with its elaborate shape and spines.

A total of 83 freshwater snail species (65 endemic) and 11 bivalve species (8 endemic) are known from the lake.<ref name=Darwall2011>Template:Cite book</ref> Among the endemic bivalves are three monotypic genera: Grandidieria burtoni, Pseudospatha tanganyicensis and Brazzaea anceyi.<ref name=Darwall2011/> 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".<ref name=Snail>Brown, D. (1994). Freshwater Snails Of Africa And Their Medical Importance. 2nd edition. Template:ISBN</ref> All the Tanganyika thalassoids, which are part of Prosobranchia, are endemic to the lake.<ref name=Snail/> 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.<ref name=LTBP/><ref name=Snail/><ref>Template:Cite journal</ref> A total of 17 freshwater snail genera are endemic to the lake, such as Hirthia, Lavigeria, Paramelania, Reymondia, Spekia, Stanleya, Tanganyicia and Tiphobia.<ref name=Snail/> 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.<ref name=Snail/> The latter is the largest Tanganyika snail and its shell is often used by small shell-dwelling cichlids.<ref>Template:Cite journal</ref>

Crustaceans are also highly diverse in Tanganyika with more than 200 species, of which more than half are endemic.<ref name=LTBP/> They include 10 species of freshwater crabs (9 Platythelphusa and Potamonautes platynotus; all endemic),<ref name=crabs>Template:Cite journal</ref> at least 11 species of small atyid shrimp (Atyella, Caridella and Limnocaridina),<ref name=Atyid>Template:Cite journal</ref> an endemic palaemonid shrimp (Macrobrachium moorei),<ref>Template:Cite iucn</ref> about 100 ostracods,<ref>Template:Cite journal</ref> including many endemics,<ref>Template:Cite journal</ref><ref>Template:Cite journal</ref> and several copepods.<ref>Template:Cite journal</ref> Among these, Limnocaridina iridinae lives inside 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).<ref>Template:Cite journal</ref><ref>Template:Cite iucn</ref>

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).<ref name=Mollusc>Segers, H.; and Martens, K; editors (2005). The Diversity of Aquatic Ecosystems. p. 46. Developments in Hydrobiology. Aquatic Biodiversity. Template:ISBN</ref> For example, the only other Rift Valley lake with endemic freshwater crabs are Lake Kivu and Lake Victoria with two species each.<ref>Cumberlidge, N.; and Meyer, K. S. (2011). A revision of the freshwater crabs of Lake Kivu, East Africa. Template:Webarchive Journal Articles. Paper 30.</ref><ref>Template:Cite journal</ref>

Other invertebratesEdit

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),<ref name=SegersMartens>Segers, H.; and Martens, K; editors (2005). The Diversity of Aquatic Ecosystems. p. 44. Developments in Hydrobiology. Aquatic Biodiversity. Template:ISBN</ref> 9 sponges (7 endemic), 6 bryozoa (2 endemic), 11 flatworms (7 endemic), 20 nematodes (7 endemic), 28 annelids (17 endemic)<ref name=LTBP/> and the small hydrozoan jellyfish Limnocnida tanganyicae.<ref>Salonen; Högmander; Langenberg; Mölsä; Sarvala; Tarvainen; and Tiirola (2012). Limnocnida tanganyicae medusae (Cnidaria: Hydrozoa): a semiautonomous microcosm in the food web of Lake Tanganyika. Hydrobiologia 690(1): 97–112.</ref>

FishingEdit

File:Fishing boat in Kagongo Ward.jpg
Fishing Boat in Kagongo Ward, Kigoma District
File:Fisherman on Lake Tanganyika.jpg
Fishermen on Lake Tanganyika

Lake Tanganyika supports a major fishery, which, depending on source, provides 25–40%<ref>{{#invoke:citation/CS1|citation |CitationClass=web }}</ref> or Template:Circa 60% of the animal protein in the diet of the people living in the region.<ref name=Jensen2016/><ref name=McGrath2016>Template:Cite news</ref>

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 the fish populations, causing significant declines.<ref name=Jensen2016/><ref name=McGrath2016/><ref name=Reilly2003/> In 2016, it was estimated that the total catch was up to 200,000 tonnes.<ref name=Jensen2016/>

HistoryEdit

File:Stanley's Lake Tanganyika.jpg
Lake Tanganyika. The black line indicates Henry Morton Stanley's route.

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.<ref>Template:Cite book</ref>

There are many methods which the native people of the area used for 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 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.<ref>Template:Cite book</ref>

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 the lake. He noted the name "Liemba" for its southern part, a word probably from the Fipa language.<ref>Template:Cite book</ref> Tanganyika means "stars" in the Luvale language.<ref>Template:Cite journal</ref>Template:Rp

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 the lake, and as a base from which to launch surprise attacks on Allied troops.<ref name=Foden>Giles Foden: Mimi and Toutou Go Forth — The Bizarre Battle for Lake Tanganyika, Penguin, 2004.</ref> 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 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.<ref name=Foden/> In order to avoid his prize ship falling into Allied hands, Zimmer scuttled the vessel on July 26, 1916. The vessel was later raised in 1924 and renamed MV Liemba.<ref name=Foden/>

ReferencesEdit

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External linksEdit

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