Lake Tanganyika

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Lake Tanganyika
Ikiyaga ca Tanganyika (Kirundi)
STS51G-034-0012 Lake Tanganyika June1985.jpg
Lake Tanganyika from space, June 1985
Africa relief location map.jpg
Red pog.svg
Lake Tanganyika
Lake Tanganyika map.png
Lake Tanganyika map
Coordinates 6°06′S29°30′E / 6.1°S 29.5°E / -6.1; 29.5
Lake type Ancient lake, Rift Valley Lake
Primary inflows Ruzizi River
Malagarasi River
Kalambo River
Primary outflows Lukuga River
Catchment area 231,000 km2 (89,000 sq mi)
Basin  countries Burundi, DR Congo, Tanzania, and Zambia
Max. length673 km (418 mi)
Max. width72 km (45 mi)
Surface area 32,900 km2 (12,700 sq mi)
Average depth570 m (1,870 ft)
Max. depth1,470 m (4,820 ft)
Water volume 18,750 km3 (4,500 cu mi)
Residence time 5500 years [1]
Shore length11,828 km (1,136 mi)
Surface elevation773 m (2,536 ft) [2]
Settlements Kigoma, Tanzania
Kalemie, the DRC
Bujumbura, Burundi
Mpulungu, Zambia
References [2]
Official nameTanganyika
Designated2 February 2007
Reference no.1671 [3]
1 Shore length is not a well-defined measure.
Lake Tanganyika eastern Shore in Kagongo Ward, Kigoma Region, Tanzania View of Kagongo Ward.jpg
Lake Tanganyika eastern Shore in Kagongo Ward, Kigoma Region, Tanzania

Lake Tanganyika ( /ˌtæŋɡənˈjkə,-ɡæn-/ TANG-gən-YEE-kə, -gan-; [4] Kirundi : Ikiyaga ca Tanganyika) is an African Great Lake. [5] It is the second-largest freshwater lake by volume and the second deepest, in both cases after Lake Baikal in Siberia. [6] [7] It is the world's longest freshwater lake. [6] 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 the Congo River system and ultimately into the Atlantic Ocean.[ citation needed ]

Contents

Geography

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 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 km2 (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 the northern basin). It holds an estimated 18,750 km3 (4,500 cu mi). [8]

The catchment area of the lake is 231,000 km2 (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 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. [9]

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. [10] The Malagarasi River, which is Tanzania's second largest river, enters the east side of Lake Tanganyika. [10] 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. [9]

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. [10] 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. [9]

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. [11]

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. [12]

Water characteristics

Clear water lake of Lake Tanganyika in Kagongo Ward, Kigoma Region, Tanzania 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 0–100 m (0–330 ft). [13] Below this, it is around 8.7, gradually decreasing to 8.3–8.5 in the deepest parts of Tanganyika. [13] 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. [13]

Surface temperatures generally range from about 24 °C (75 °F) in the southern part of the lake in early August to 28–29 °C (82–84 °F) in the late rainy season in March—April. [14] 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). [15] The water has gradually warmed since the 19th century and this has accelerated with global warming since the 1950s. [16]

The lake is stratified and seasonal mixing generally does not extend beyond depths of 150 m (490 ft). [14] 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. [17] As a consequence of the stratification, the deep sections contain "fossil water". [18] 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 100 m (330 ft) in the northern part of the lake and 240–250 m (790–820 ft) in the south. [19] [20] The oxygen-devoid deepest sections contain high levels of toxic hydrogen sulphide and are essentially lifeless, [6] except for bacteria. [13] [21]

Biology

Reptiles

Nile crocodiles swimming CrocoLoco-swimming-01.jpg
Nile crocodiles swimming

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). [22] 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. [22] [23]

Cichlid fish

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. 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.
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 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

The lake holds at least 250 species of cichlid fish [27] and undescribed species remain. [28] Almost all (98%) of the Tanganyika cichlids are endemic to the lake and it is thus an important biological resource for the study of speciation in evolution. [29] [30] Some of the endemics do occur slightly into the upper Lukuga River, Lake Tanganyika's outflow, but further spread into the Congo River basin is prevented by physics (Lukuga has fast-flowing sections with many rapids and waterfalls) and chemistry (Tanganyika's water is alkaline, while the Congo's generally is acidic). [9] The cichlids of the African Great Lakes, including Tanganyika, represent the most diverse extent of adaptive radiation in vertebrates. [31]

Although Tanganyika has far fewer cichlid species than Lakes Malawi and Victoria which both have experienced relatively recent explosive species radiations (resulting in many closely related species), [32] its cichlids are the most morphologically and genetically diverse. [31] [33] This is linked to the high age of Tanganyika, as it is far older than the other lakes. [34] Tanganyika has the largest number of endemic cichlid genera of all African lakes. [31] 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) and another three have species in the lake (Haplochromini, Tilapiini and Tylochromini). [35] Others have proposed splitting the Tanganyika cichlids into as many as 12–16 tribes (in addition to previous mentioned, Bathybatini, Benthochromini, Boulengerochromini, Cyphotilapiini, Eretmodini, Greenwoodochromini, Perissodini and Trematocarini). [31]

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). [36] Trematocara species have exceptionally been found at more than 300 m (980 ft), which is deeper than any other cichlid in the world. [37] Some of the deep-water cichlids (e.g., Bathybates , Gnathochromis , Hemibates and Xenochromis ) have been caught in places virtually devoid of oxygen, but how they are able to survive there is unclear. [20] Tanganyika cichlids are generally benthic (found at or near the bottom) and/or coastal. [38] No Tanganyika cichlids are truly pelagic and offshore, except for some of the piscivorous Bathybates. [36] Two of these, B. fasciatus and B. leo , mainly feed on Tanganyika sardines. [36] [20] Tanganyika cichlids differ extensively in ecology and include species that are herbivores, detritivores, planktivores, insectivores, molluscivores, scavengers, scale-eaters and piscivores. [28] These dietary specializations, however, have been shown to be flexible. 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. [39] Their breeding behavior fall into two main groups, the substrate spawners (often in caves or rock crevices) and the mouthbrooders. [40] Among the endemic species are two of the world's smallest cichlids, Neolamprologus multifasciatus and N. similis (both shell dwellers) at up to 4–5 cm (1.6–2.0 in), [41] [42] and one of the largest, the giant cichlid (Boulengerochromis microlepis) at up to 90 cm (3.0 ft). [28] [43]

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. [40] Recreating a Lake Tanganyika biotope to host those cichlids in a habitat similar to their natural environment is also popular in the aquarium hobby. [40] [44]

Other fish

The Tanganyika killifish (Lamprichthys tanganicanus) is the only member of its genus Lamprichthys tanganicanus.JPG
The Tanganyika killifish (Lamprichthys tanganicanus) is the only member of its genus

Lake Tanganyika is home to more than 80 species of non-cichlid fish and about 60% of these are endemic. [19] [27] [63] [64]

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). [38] 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. [38] The four lates, all endemic to Tanganyika, have been overfished and larger individuals are rare today. [38]

Among the more unusual fish in the lake are the endemic, facultatively brood parasitic "cuckoo catfish", including at least Synodontis grandiops [65] and S. multipunctatus . [19] [40] 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. [19] [66] 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. [19] [40] Six catfish genera are entirely restricted to the lake basin: Bathybagrus , Dinotopterus , Lophiobagrus , Phyllonemus , Pseudotanganikallabes and Tanganikallabes . [53] [67] 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; [53] in the latter habitat they are the primary predators and scavengers. [20] A unique evolutionary radiation in the lake is the 15 species of Mastacembelus spiny eels, all but one endemic to its basin. [63] [68] 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. [68]

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). [53]

Molluscs and crustaceans

The shell of the endemic thalassoid freshwater snail Tiphobia horei with its elaborate shape and spines. 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. [69] Among the endemic bivalves are three monotypic genera: Grandidieria burtoni , Pseudospatha tanganyicensis and Brazzaea anceyi . [69] 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". [70] All the Tanganyika thalassoids, which are part of Prosobranchia, are endemic to the lake. [70] 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. [27] [70] [71] A total of 17 freshwater snail genera are endemic to the lake, such as Hirthia , Lavigeria , Paramelania , Reymondia , Spekia , Stanleya , Tanganyicia and Tiphobia . [70] 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 . [70] The latter is the largest Tanganyika snail and its shell is often used by small shell-dwelling cichlids. [72]

Crustaceans are also highly diverse in Tanganyika with more than 200 species, of which more than half are endemic. [27] They include 10 species of freshwater crabs (9 Platythelphusa and Potamonautes platynotus ; all endemic), [73] at least 11 species of small atyid shrimp ( Atyella , Caridella and Limnocaridina ), [74] an endemic palaemonid shrimp ( Macrobrachium moorei), [75] about 100 ostracods, [76] including many endemics, [77] [78] and several copepods. [79] 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). [80] [81]

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). [82] For example, the only other Rift Valley lake with endemic freshwater crabs are Lake Kivu and Lake Victoria with two species each. [83] [84]

Other invertebrates

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), [85] 9 sponges (7 endemic), 6 bryozoa (2 endemic), 11 flatworms (7 endemic), 20 nematodes (7 endemic), 28 annelids (17 endemic) [27] and the small hydrozoan jellyfish Limnocnida tanganyicae. [86]

Fishing

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

Lake Tanganyika supports a major fishery, which, depending on source, provides 25–40% [87] or c. 60% of the animal protein in the diet of the people living in the region. [16] [88]

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. [16] [88] [15] In 2016, it was estimated that the total catch was up to 200,000 tonnes. [16]

History

Lake Tanganyika. The black line indicates Henry Morton Stanley's route. 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. [89]

There are many methods in which the native people of 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 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. [90]

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. [91] Tanganyika means "stars" in the Luvale language. [92] :523

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. [93] 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. [93] 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. [93]

Related Research Articles

<span class="mw-page-title-main">Cichlid</span> Family of fishes

Cichlids are fish from 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 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 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.

<span class="mw-page-title-main">Lake Malawi</span> Lake in Africa

Lake Malawi, also known as Lake Nyasa in Tanzania and Lago Niassa in Mozambique, is an African Great Lake and the southernmost lake in the East African Rift system, located between Malawi, Mozambique and Tanzania.

<span class="mw-page-title-main">Lake Victoria</span> Lake in East-central Africa

Lake Victoria is one of the African Great Lakes. With a surface area of approximately 59,947 km2 (23,146 sq mi), Lake Victoria is Africa's largest lake by area, the world's largest tropical lake, and the world's second-largest fresh water lake by surface area after Lake Superior in North America. In terms of volume, Lake Victoria is the world's ninth-largest continental lake, containing about 2,424 km3 (1.965×109 acre⋅ft) of water. Lake Victoria occupies a shallow depression in Africa. The lake has an average depth of 40 m (130 ft) and a maximum depth of 80–81 m (262–266 ft). Its catchment area covers 169,858 km2 (65,583 sq mi). The lake has a shoreline of 7,142 km (4,438 mi) when digitized at the 1:25,000 level, with islands constituting 3.7% of this length.

<span class="mw-page-title-main">Lake Kivu</span> Meromictic lake in the East African Rift valley

Lake Kivu is one of the African Great Lakes. It lies on the border between the Democratic Republic of the Congo and Rwanda, and is in the Albertine Rift, the western branch of the East African Rift. Lake Kivu empties into the Ruzizi River, which flows southwards into Lake Tanganyika. In 1894, German explorer and officer Gustav Adolf von Götzen was the first European to discover the lake.

<i>Lamprologus ocellatus</i> Species of fish

Lamprologus ocellatus is a species of shell dwelling cichlid endemic to Lake Tanganyika. It is a popular aquarium fish due to its small size, appearance, and intelligence.

<i>Synodontis multipunctatus</i> Species of fish

Synodontis multipunctatus, also known as the cuckoo catfish, cuckoo squeaker, or multipunk, is a small catfish from Lake Tanganyika, one of the lakes in the Great Rift Valley system in Africa. It is a brood parasite upon mouthbrooding cichlids. This species grows to a length of 27.5 centimetres (10.8 in) TL. This species is a minor component of local commercial fisheries.

<i>Synodontis</i> Genus of fishes

Synodontis is the largest genus of mochokid catfishes. It is the biggest genus within the 10 genera and 190 different species in the family Mochokidae. Synodontis has over 131 different species within the genus. Synodontis are also known as squeakers, due to their ability to make stridulatory sounds through their pectoral fin spines when handled or disturbed. Synodontis make a sound that sounds like squeaking by rubbing their spines together. They do this when they have been frightened or when they become angry. Synodontis may also squeak when they are taken out of the water. These catfish are small- to medium-sized fish with many species exhibiting attractive spotted markings. Some species are also known for naturally swimming belly-up, earning the name upside-down catfish. Some of these species are Synodontis contractus and Synodontis nigriventris. While some of these species are known to swim upside down, another species, Synodontis multipunctatus, is a brood parasitic cuckoo catfish,there are two other species Synodontis petricola and Synodontis grandiops are also called brood parasitic cuckoo catfish.

<span class="mw-page-title-main">Lepidophagy</span> Fish eating scales of other fish

Lepidophagy is a specialised feeding behaviour in fish that involves eating the scales of other fish. Lepidophagy is widespread, having evolved independently in at least five freshwater families and seven marine families. A related feeding behavior among fish is pterygophagy: feeding on the fins of other fish.

<i>Altolamprologus compressiceps</i> Species of fish

Altolamprologus compressiceps is a species of fish in the family Cichlidae. It is endemic to the shallow rocky areas of Lake Tanganyika. It is not considered threatened by the IUCN.

<span class="mw-page-title-main">Convict julie</span> Species of fish

The convict julie is a cichlid species in the subfamily Pseudocrenilabrinae family endemic to Lake Tanganyika. Hence it is found in Burundi, the Democratic Republic of the Congo, Tanzania, and Zambia. The fish is named after Charles Tate Regan.

<i>Perissodus</i> Genus of fishes

Perissodus is a genus of cichlids endemic to Lake Tanganyika in Africa. They feed on scales.

Plecodus elaviae is a species of cichlid endemic to Lake Tanganyika. This schooling species is a scale-eater, plucking scales from other fishes. Both parents care for the offspring in this mouthbrooder. This fish can reach a length of 32 centimetres (13 in) TL.

<i>Plecodus</i> Genus of fishes

Plecodus is a genus of cichlids endemic to Lake Tanganyika in Africa. They feed on scales.

<span class="mw-page-title-main">Spotfin goby cichlid</span> Species of fish

The spotfin goby cichlid is an African species of cichlid endemic to Lake Tanganyika where it is only known from the northern end of the lake. They live amongst pebbles in the surf-zone. This species can reach a length of 7 centimetres (2.8 in) TL. This species can also be found in the aquarium trade. Although presently considered the only species in the genus, another undescribed species is known from the Lukuga River.

<i>Neothauma</i> Genus of gastropods

Neothauma is a genus of freshwater snail with a gill and an operculum, an aquatic gastropod mollusc in the subfamily Bellamyinae of the family Viviparidae.

<i>Synodontis petricola</i> Species of fish

Synodontis petricola, known as the cuckoo catfish, or the pygmy leopard catfish, is a species of upside-down catfish endemic to Burundi, Zambia, the Democratic Republic of the Congo, and Tanzania where it is only known from Lake Tanganyika. It was first described by Belgian ichthyologist Hubert Matthes in 1959. The species name "petricola" is derived from a combination of the Latin petra, meaning stone or rock, and the Latin cola, meaning inhabitant. This refers to the rocky environment where this species is found.

<span class="mw-page-title-main">Eretmodini</span> Tribe of fishes

Eretmodini is a tribe of African cichlids. It contains five species of freshwater fish endemic to Lake Tanganyika. They are small fish with reduced swim bladders that are found near the bottom in the turbulent, coastal surf zone. They are mouthbrooders.

<span class="mw-page-title-main">Perissodini</span> Tribe of fishes

Perissodini is a tribe of African cichlids, containing three genera of freshwater fish found only in Lake Tanganyika.

Synodontis dorsomaculatus is a species of upside-down catfish that is endemic to the Democratic Republic of the Congo where it occurs in the Lukusi and upper Lualaba Rivers and the Upemba lakes. It was first described by Max Poll in 1971. The original specimens were obtained in Kadia, Kisale, Katanga, in what is now the Democratic Republic of the Congo. The species name dorsomaculatus is derived from the word dorso, meaning "back", and the word maculatus, meaning "spotted", a reference to the dark round spots on the fish's back.

<i>Mastacembelus ellipsifer</i> Species of fish

Mastacembelus ellipsifer is a species of spiny eel that is endemic to Lake Tanganyika in Africa and sometimes kept in aquariums. Although sometimes called the Tanganyikan spiny eel, it is only one of fifteen spiny eel species in the Tanganyikan basin.

References

  1. Yohannes, Okbazghi (2008). Water resources and inter-riparian relations in the Nile basin. SUNY Press. p. 127.
  2. 1 2 "LAKE TANGANYIKA". www.ilec.or.jp. Archived from the original on 28 March 2008. Retrieved 14 March 2008.
  3. "Tanganyika". Ramsar Sites Information Service. Archived from the original on 27 April 2018. Retrieved 25 April 2018.
  4. Wells, John C. (2008). Longman Pronunciation Dictionary (3rd ed.). Longman. ISBN   978-1-4058-8118-0.
  5. Francis., Burton, Richard (1965). Burton and Lake Tanganyika. East African Literature Bureau. OCLC   180480726.{{cite book}}: CS1 maint: multiple names: authors list (link)
  6. 1 2 3 "Lake Tanganyika". www.zambiatourism.com. Archived from the original on 22 April 2015. Retrieved 14 March 2008.
  7. Lewis, R. (16 May 2010). "Brown Geologists Show Unprecedented Warming in Lake Tanganyika". Brown University. Archived from the original on 26 March 2017. Retrieved 25 March 2017.
  8. "Datbase Summary: Lake Tanganyika". ilec.or.jp. Japan: International Lake Environment Committee Foundation. Archived from the original on 10 November 1999. Retrieved 14 March 2008.
  9. 1 2 3 4 Kullander, S.O.; Roberts, T.R. (2011). "Out of Lake Tanganyika: endemic lake fishes inhabit rapids of the Lukuga River". Ichthyol. Explor. Freshwaters. 22 (4): 355–76.
  10. 1 2 3 Scheffel, Richard L.; Wernet, Susan J., eds. (1980). Natural Wonders of the World. US: Reader's Digest Association, Inc. pp. 366–67. ISBN   978-0-89577-087-5.
  11. Lévêqu, Christian (1997). Biodiversity Dynamics and Conservation: The Freshwater Fish of Tropical Africa. Cambridge University Press. p. 110.
  12. Cohen; Soreghan; Scholz (1993). "Estimating the age of formation of lakes: An example from Lake Tanganyika, East African Rift system". Geosciences. 21 (6): 511–514. Bibcode:1993Geo....21..511C. doi:10.1130/0091-7613(1993)021<0511:ETAOFO>2.3.CO;2.
  13. 1 2 3 4 De, Wever; Muylaert; der Gucht, Van; Pirlot; Cocquyt; Descy; Plisnier; Vyverman (2005). "Bacterial Community Composition in Lake Tanganyika: Vertical and Horizontal Heterogeneity". Appl Environ Microbiol. 71 (9): 5029–5037. Bibcode:2005ApEnM..71.5029D. doi:10.1128/AEM.71.9.5029-5037.2005. PMC   1214687 . PMID   16151083.
  14. 1 2 Edmond; Stallard; Craigh; Weiss; Coulter (1993). "Nutrient chemistry of the water column of Lake Tanganyika". Limnol. Oceanogr. 38 (4): 725–738. Bibcode:1993LimOc..38..725E. doi: 10.4319/lo.1993.38.4.0725 .
  15. 1 2 O'Reilly, Catherine M.; Alin, Simone R.; Plisnier, Pierre-Denis; Cohen, Andrew S.; Mckee, Brent A. (14 August 2003). "Climate change decreases aquatic ecosystem productivity of Lake Tanganyika, Africa". Nature. 424 (6950): 766–768. Bibcode:2003Natur.424..766O. doi:10.1038/nature01833. PMID   12917682. S2CID   1637315.
  16. 1 2 3 4 Jensen, M.R. (8 August 2016). "Lake Tanganyika Fisheries Declining From Global Warming". University of Arizona. Archived from the original on 6 March 2018. Retrieved 5 March 2018.
  17. Lowe-McConnell, R.H. (2003). "Recent research in the African Great Lakes: Fisheries, biodiversity and cichlid evolution". Freshwater Forum. 20 (1): 4–64.
  18. Hutter; Yongqi; and Chubarenko (2011). Physics of Lakes, volume 1: Foundation of the Mathematical and Physical Background. P. 11. ISBN   978-3-642-15178-1.
  19. 1 2 3 4 5 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.
  20. 1 2 3 4 Lowe-McConnell, R.H. (1987). Ecological Studies in Tropical Fish Communities. ISBN   0-521-28064-8.
  21. Ryan, Emily; Todd, Jonathan A.; McGlue, Michael; Kimirei, Ismael; Soreghan, Michael (2017). "Variation in Taphonomic Character of Shell Beds in Lake Tanganyika, Africa: Paleoenvironmental and Stratigraphic Implications of Shell Beds in Lakes". Geological Society of America Abstracts with Programs. Geological Society of America. doi:10.1130/abs/2017am-304608.{{cite journal}}: Cite journal requires |journal= (help)
  22. 1 2 Spawls, Howell, Drewes, and Ashe (2002). A Field Guide to the Reptiles of East Africa. Academic Press, London. ISBN   0-12-656470-1.
  23. O'Shea, Mark; Mcintyre, P.B. (2005). "Boulengerina annulata stormsi (Storm's water cobra). Attempted predation". Herpetological Review. 36 (2): 189. Archived from the original on 2 December 2017. Retrieved 11 March 2023.
  24. Dierkesa; Taborskya; Kohler (1999). "Reproductive parasitism of broodcare helpers in a cooperatively breeding fish". Behavioral Ecology. 10 (5): 510–515. doi: 10.1093/beheco/10.5.510 .
  25. Balshine-Earn; Lotem (1998). "Individual recognition in a cooperatively breeding cichlid : Evidence from video playback experiments". Behaviour. 135 (3): 369–386. doi:10.1163/156853998793066221.
  26. Wernera; Balshineb; Leachc; Lotem (2003). "Helping opportunities and space segregation in cooperatively breeding cichlids". Behavioral Ecology. 14 (6): 749–756. doi: 10.1093/beheco/arg067 .
  27. 1 2 3 4 5 Kelly West (28 February 2001). "Results and Experiences of the UNDP/GEF Conservation Initiative (RAF/92/G32) in Burundi, D.R. Congo, Tanzania, and Zambia". IW:LEARN | Documents. Archived from the original on 11 March 2023. Retrieved 11 March 2023.
  28. 1 2 3 Craig Mortiff (24 December 2009). "Lake Tanganyika and its Diverse Cichlids". Cichlid Fish Forum. Archived from the original on 11 March 2023. Retrieved 11 March 2023.
  29. Takahashi, T.; Hori, M. (2012). "Genetic and Morphological Evidence Implies Existence of Two Sympatric Species in Cyathopharynx furcifer (Teleostei: Cichlidae) from Lake Tanganyika". International Journal of Evolutionary Biology. 2012: 980879. doi: 10.1155/2012/980879 . PMC   3363988 . PMID   22675655.
  30. Kornfield, Ivy; Smith, Peter A (2000). "African Cichlid Fishes: Model Systems for Evolutionary Biology". Annual Review of Ecology and Systematics. 31: 163–196. doi:10.1146/annurev.ecolsys.31.1.163.
  31. 1 2 3 4 Meyer, Britta; Matchiner, Michael; Salburger, Walter (25 November 2013). "A tribal level phylogeny of Lake Tanganyika cichlid fishes based on a genomic multi-marker approach". Molecular Phylogenetics and Evolution. 83: 56–71. doi:10.1016/j.ympev.2014.10.009. PMC   4334724 . PMID   25433288.
  32. Turner, Seehausen; Knight, Allender; Robinson (2001). "How many species of cichlid fishes are there in African lakes?". Molecular Ecology. 10 (3): 793–806. Bibcode:2001MolEc..10..793T. doi: 10.1046/j.1365-294x.2001.01200.x . PMID   11298988. S2CID   12925712.
  33. Seehausen, O. (2015). "Process and pattern in cichlid radiations – inferences for understanding unusually high rates of evolutionary diversification". New Phytologist. 207 (2): 304–312. doi: 10.1111/nph.13450 . PMID   25983053.
  34. Nishida, M (1991). "Lake Tanganyika as an evolutionary reservoir of old lineages of East African cichlid fishes: Inferences from allozyme data". Experientia. 47 (9): 974–979. doi:10.1007/bf01929896. S2CID   37599331.
  35. Sparks; Smith (2004). "Phylogeny and biogeography of cichlid fishes (Teleostei: Perciformes: Cichlidae)". Cladistics. 20 (6): 501–517. CiteSeerX   10.1.1.595.2118 . doi:10.1111/j.1096-0031.2004.00038.x. PMID   34892958. S2CID   36086310.
  36. 1 2 3 4 Kirchberger; Sefc; Sturmbauer; Koblmuller (2012). "Evolutionary History of Lake Tanganyika's Predatory Deepwater Cichlids". International Journal of Evolutionary Biology. 2012: 716209. doi: 10.1155/2012/716209 . PMC   3362839 . PMID   22675652.
  37. Loiselle, Paul (1994). The Cichlid Aquarium, p. 304. Tetra Press, Germany. ISBN   978-1564651464.
  38. 1 2 3 4 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. ISBN   978-0792360179
  39. Golcher-Benavides J, Wagner CE (2019). "Playing out Liem's Paradox: Opportunistic Piscivory across Lake Tanganyikan Cichlids". The American Naturalist. 194 (2): 260–267. doi:10.1086/704169. PMID   31318283. S2CID   155438716.
  40. 1 2 3 4 5 Schliewen, U. (1992). Aquarium Fish. Barron's Educational Series. ISBN   978-0812013504.
  41. Froese, Rainer; Pauly, Daniel (eds.). "Neolamprologus multifasciatus". FishBase . March 2017 version.
  42. Froese, Rainer; Pauly, Daniel (eds.). "Neolamprologus similis". FishBase . March 2017 version.
  43. 1 2 "The 10 biggest cichlids". Practical Fishkeeping. 13 June 2016. Archived from the original on 20 March 2017. Retrieved 17 March 2017.
  44. "tanganyika biotope aquarium". Aquariums Life. 10 February 2010. Archived from the original on 2 March 2012. Retrieved 3 February 2014.{{cite web}}: CS1 maint: unfit URL (link)
  45. Meyer; Matchiner; Salburger (2015). "Lake Tanganyika—A 'Melting Pot' of Ancient and Young Cichlid Lineages (Teleostei: Cichlidae)?". PLOS ONE. 10 (7): e0125043. Bibcode:2015PLoSO..1025043W. doi: 10.1371/journal.pone.0125043 . PMC   4415804 . PMID   25928886.
  46. 1 2 Weiss; Cotterill; Schliewen (2015). "A tribal level phylogeny of Lake Tanganyika cichlid fishes based on a genomic multi-marker approach". Molecular Phylogenetics and Evolution. 83: 56–71. doi:10.1016/j.ympev.2014.10.009. PMC   4334724 . PMID   25433288.
  47. Takahashi, T. (2008). "Description of a new cichlid fish species of the genus Benthochromis (Perciformes: Cichlidae) from Lake Tanganyika". Journal of Fish Biology. 72 (3): 603–613. Bibcode:2008JFBio..72..603T. doi:10.1111/j.1095-8649.2007.01727.x.
  48. Takahashi, T.; Nakaya, K. (2003). "New species of Cyphotilapia (Perciformes: Cichlidae) from Lake Tanganyika, Africa". Copeia. 2003 (4): 824–832. doi:10.1643/ia03-148.1. S2CID   83854866.
  49. Bigirimana, C. (2006). "Cyprichromis microlepidotus". IUCN Red List of Threatened Species . 2006: e.T60487A12363286. doi: 10.2305/IUCN.UK.2006.RLTS.T60487A12363286.en . Retrieved 13 November 2021.
  50. 1 2 Smith, M.P. (1998). Lake Tanganyikan Cichlids, pp. 9–10. ISBN   0-7641-0615-5
  51. "Ophthalmotilapia nasuta". Seriously Fish. Archived from the original on 11 March 2023. Retrieved 11 March 2023.
  52. Froese, Rainer; Pauly, Daniel (eds.). "Eretmodus cyanostictus". FishBase . April 2017 version.
  53. 1 2 3 4 "Species in the Tanganyika". www.fishbase.se (table). Archived from the original on 11 March 2023. Retrieved 11 March 2023.
  54. Lowe-McConnell, R (2009). "Fisheries and cichlid evolution in the African Great Lakes: progress and problems". Freshwater Reviews. 2 (2): 131–151. doi:10.1608/frj-2.2.2. S2CID   54011001.
  55. "Julidochromis marlieri (Marlier's Julie)". Seriously Fish. Archived from the original on 11 March 2023. Retrieved 11 March 2023.
  56. Bigirimana, C. (2006). "Gnathochromis permaxillaris". IUCN Red List of Threatened Species . 2006: e.T60493A12364587. doi: 10.2305/IUCN.UK.2006.RLTS.T60493A12364587.en . Retrieved 13 November 2021.
  57. Stewart, T.A.; Albertson, R.C. (2010). "Evolution of a unique predatory feeding apparatus: functional anatomy, development and a genetic locus for jaw laterality in Lake Tanganyika scale-eating cichlids". BMC Biology. 8 (1): 8. doi: 10.1186/1741-7007-8-8 . PMC   2828976 . PMID   20102595.
  58. Ntakimazi, G. (2006). "Oreochromis tanganicae". IUCN Red List of Threatened Species . 2006: e.T60625A12387918. doi: 10.2305/IUCN.UK.2006.RLTS.T60625A12387918.en . Retrieved 13 November 2021.
  59. Begon, M.; and A.H. Fitter (1995). Advances in Ecological Research, vol. 26, p. 203. ISBN   0-12-013926-X
  60. Salzburger; Niederstätter; Brandstätter; Berger; Parson; Snoeks; Sturmbauer (2006). "Colour-assortative mating among populations of Tropheus moorii, a cichlid fish from Lake Tanganyika, East Africa". Proc Biol Sci. 273 (1584): 257–266. doi:10.1098/rspb.2005.3321. PMC   1560039 . PMID   16543167.
  61. Robert Toman (2017). "Tropheus Genus Evolution". Cichlid World. Archived from the original on 19 September 2021. Retrieved 11 March 2023.
  62. Froese, Rainer; Pauly, Daniel (eds.). "Species in genus Lamprichthys". FishBase . March 2017 version.
  63. 1 2 Brown; Britz; Bills; Rüber; Day (2011). "Pectoral fin loss in the Mastacembelidae: a new species from Lake Tanganyika". Journal of Zoology. 284 (4): 286–293. doi:10.1111/j.1469-7998.2011.00804.x.
  64. Wright, J.J.; Bailey, R.M. (2012). "Systematic revision of the formerly monotypic genus Tanganikallabes (Siluriformes: Clariidae)" (PDF). Zoological Journal of the Linnean Society. 165 (1): 121–142. doi: 10.1111/j.1096-3642.2011.00789.x .
  65. "Synodontis grandiops • Mochokidae". www.planetcatfish.com. 2020. Archived from the original on 1 April 2017. Retrieved 11 March 2023.
  66. "Synodontis lucipinnis • Mochokidae". www.planetcatfish.com. 2023. Archived from the original on 1 April 2017. Retrieved 11 March 2023.
    "Synodontis petricola • Mochokidae". www.planetcatfish.com. 2023. Archived from the original on 1 April 2017. Retrieved 11 March 2023.
  67. Wright, J.J. (2017). "A new diminutive genus and species of catfish from Lake Tanganyika (Siluriformes: Clariidae)". J Fish Biol. 91 (3): 789–805. Bibcode:2017JFBio..91..789W. doi:10.1111/jfb.13374. PMID   28744868.
  68. 1 2 Brown; Rüber; Bills; Day (2010). "Mastacembelid eels support Lake Tanganyika as an evolutionary hotspot of diversification". BMC Evolutionary Biology. 10: 188. doi: 10.1186/1471-2148-10-188 . PMC   2903574 . PMID   20565906.
  69. 1 2 Seddon, M.; Appleton, C.; Van Damme, D.; Graf, D. (2011). Darwall, W.; Smith, K.; Allen, D.; Holland, R.; Harrison, I.; Brooks, E. (eds.). Freshwater molluscs of Africa: diversity, distribution, and conservation. IUCN. pp. 92–119. ISBN   978-2831713458.{{cite book}}: |journal= ignored (help)
  70. 1 2 3 4 5 Brown, D. (1994). Freshwater Snails Of Africa And Their Medical Importance. 2nd edition. ISBN   0-7484-0026-5
  71. West, K.; Cohen, A. (1996). "Shell microstructure of gastropods from Lake Tanganyika, Africa: adaptation, convergent evolution, and escalation". Evolution. 50 (2): 672–682. doi:10.2307/2410840. JSTOR   2410840. PMID   28568935.
  72. Koblmüller; Duftner; Sefc; Aibara; Stipacek; Blanc; Egger; Sturmbauer (2007). "Reticulate phylogeny of gastropod-shell-breeding cichlids from Lake Tanganyika — the result of repeated introgressive hybridization". BMC Evolutionary Biology. 7: 7. doi: 10.1186/1471-2148-7-7 . PMC   1790888 . PMID   17254340.
  73. Marijnissen; Michel; Daniels; Erpenbeck; Menken; Schram (2006). "Molecular evidence for recent divergence of Lake Tanganyika endemic crabs (Decapoda: Platythelphusidae)". Molecular Phylogenetics and Evolution. 40 (2): 628–634. doi:10.1016/j.ympev.2006.03.025. PMID   16647274.
  74. Fryer, G (2006). "Evolution in ancient lakes: radiation of Tanganyikan atyid prawns and speciation of pelagic cichlid fishes in Lake Malawi". Hydrobiologia. 568 (1): 131–142. doi:10.1007/s10750-006-0322-x. S2CID   44127332.
  75. De Grave, S. (2013). "Macrobrachium moorei". IUCN Red List of Threatened Species . 2013: e.T196882A2477768. doi: 10.2305/IUCN.UK.2013-1.RLTS.T196882A2477768.en . Retrieved 13 November 2021.
  76. Martens; Schön; Meisch; Horne (2008). "Global diversity of ostracods (Ostracoda, Crustacea) in freshwater". Hydrobiologia. 595: 185–193. doi:10.1007/s10750-007-9245-4. S2CID   207150861.
  77. Gitter, F.; Gross, M.; Piller, W.E. (2015). "Sub-Decadal Resolution in Sediments of Late Miocene Lake Pannon Reveals Speciation of Cyprideis (Crustacea, Ostracoda)". PLOS ONE. 10 (4): e0109360. Bibcode:2015PLoSO..1009360G. doi: 10.1371/journal.pone.0109360 . PMC   4406499 . PMID   25902063.
  78. Schön, I.; Martens, K. (2012). "Molecular analyses of ostracod flocks from Lake Baikal and Lake Tanganyika". Hydrobiologia. 682 (1): 91–110. doi:10.1007/s10750-011-0935-6. S2CID   14831643.
  79. Cirhuza, D.M.; Plisnier, P.-D. (2016). "Composition and seasonal variations in abundance of Copepod (Crustacea) populations from the northern part of Lake Tanganyika". Aquatic Ecosystem Health & Management. 19 (4): 401–410. doi:10.1080/14634988.2016.1251277. hdl:2268/248670. S2CID   90502032. Archived from the original on 22 July 2021. Retrieved 30 August 2020.
  80. De Grave, S.; Cai, Y.; Amnker, A. (2008). "Global diversity of shrimps (Crustacea: Decapoda: Caridea) in freshwater". Hydrobiologia. 595: 287–293. doi:10.1007/s10750-007-9024-2. S2CID   22945163.
  81. De Grave, S. (2013). "Limnocaridina iridinae". IUCN Red List of Threatened Species . 2013: e.T198058A2510158. doi: 10.2305/IUCN.UK.2013-1.RLTS.T198058A2510158.en . Retrieved 13 November 2021.
  82. Segers, H.; and Martens, K; editors (2005). The Diversity of Aquatic Ecosystems. p. 46. Developments in Hydrobiology. Aquatic Biodiversity. ISBN   1-4020-3745-7
  83. Cumberlidge, N.; and Meyer, K. S. (2011). A revision of the freshwater crabs of Lake Kivu, East Africa. Archived 25 July 2014 at the Wayback Machine Journal Articles. Paper 30.
  84. Cumberlidge, N.; Clark, P.F. (2017). "Description of three new species of Potamonautes MacLeay, 1838 from the Lake Victoria region in southern Uganda, East Africa (Brachyura: Potamoidea: Potamonautidae)". European Journal of Taxonomy (371): 1–19. doi: 10.5852/ejt.2017.371 . hdl: 10141/622400 .
  85. Segers, H.; and Martens, K; editors (2005). The Diversity of Aquatic Ecosystems. p. 44. Developments in Hydrobiology. Aquatic Biodiversity. ISBN   1-4020-3745-7
  86. 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.
  87. "Global warming is killing off tropical lake fish – Study of Lake Tanganyika". www.mongabay.com. Archived from the original on 29 June 2015. Retrieved 14 March 2008.
  88. 1 2 McGrath, M. (8 August 2016). "Decline of fishing in Lake Tanganyika 'due to warming'". BBC. Archived from the original on 28 April 2018. Retrieved 5 March 2018.
  89. East African Ecosystems and Their Conservation. New York: Oxford University Press.
  90. Lake Tanganyika and Its Life. Oxford Press. 1991.
  91. David Livingstone (2008). The Last Journals of David Livingstone in Central Africa from 1865 to His Death. Vol. 1. BiblioBazaar. p. 338. ISBN   978-0-554-26021-1.
  92. Daniel J. Crowley (Summer 1966). "An African Aesthetic". The Journal of Aesthetics and Art Criticism. 24 (4). Wiley: 519–524. doi:10.2307/428776. JSTOR   428776.
  93. 1 2 3 Giles Foden: Mimi and Toutou Go Forth — The Bizarre Battle for Lake Tanganyika, Penguin, 2004.