Editing Lake Tanganyika (section)

===Cichlid fishes===
[[File:Murky waters.jpg|thumb|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|thumb|One of the many Tanganyika cichlids is ''[[Neolamprologus brichardi]]''. The complex behaviors of this species and its close relative ''[[Neolamprologus pulcher|N. pulcher]]'' have been studied in detail<ref>{{cite journal |last1=Dierkesa |last2=Taborskya |last3=Kohler |year=1999 |title=Reproductive parasitism of broodcare helpers in a cooperatively breeding fish |journal=Behavioral Ecology |volume=10 |issue=5 |pages=510–515 |doi=10.1093/beheco/10.5.510 |doi-access=free}}</ref><ref>{{cite journal |last1=Balshine-Earn |last2=Lotem |year=1998 |title=Individual recognition in a cooperatively breeding cichlid : Evidence from video playback experiments |journal=Behaviour |volume=135 |issue=3 |pages=369–386 |doi=10.1163/156853998793066221}}</ref><ref>{{cite journal |last1=Wernera |last2=Balshineb |last3=Leachc |last4=Lotem |year=2003 |title=Helping opportunities and space segregation in cooperatively breeding cichlids |doi=10.1093/beheco/arg067 |journal=Behavioral Ecology |volume=14 |issue=6 |pages=749–756 |doi-access=free}}</ref>]]
Lake Tanganyika is home to at least 250 [[Endemism|endemic]] species of [[cichlid|cichlids]],<ref name=LTBP>{{Cite web |website=IW:LEARN {{pipe}} Documents |author1=Kelly West |title=Results and Experiences of the UNDP/GEF Conservation Initiative (RAF/92/G32) in Burundi, D.R. Congo, Tanzania, and Zambia |url=https://iwlearn.net/documents/6644 |access-date=2023-03-11 |date=28 February 2001 |archive-date=11 March 2023 |archive-url=https://web.archive.org/web/20230311162707/https://iwlearn.net/documents/6644 |url-status=live }}</ref> and [[Undescribed taxon|undescribed species]] likely remain to be discovered.<ref name=MortiffTangDiv>{{Cite web |title=Lake Tanganyika and its Diverse Cichlids |url=https://www.cichlid-forum.com/threads/lake-tanganyika-and-its-diverse-cichlids.456196/ |access-date=2023-03-11 |website=Cichlid Fish Forum |date=24 December 2009 |author1=Craig Mortiff |archive-date=11 March 2023 |archive-url=https://web.archive.org/web/20230311162711/https://www.cichlid-forum.com/threads/lake-tanganyika-and-its-diverse-cichlids.456196/ |url-status=live }}</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>{{cite journal |last1=Takahashi |first1=T. |last2=Hori |first2=M. |year=2012 |title=Genetic and Morphological Evidence Implies Existence of Two Sympatric Species in Cyathopharynx furcifer (Teleostei: Cichlidae) from Lake Tanganyika |journal=International Journal of Evolutionary Biology |volume=2012 |page=980879 |doi=10.1155/2012/980879 |pmid=22675655 |pmc=3363988 |doi-access=free}}</ref><ref>{{cite journal |last1=Kornfield |first1=Ivy |last2=Smith |first2=Peter A |year=2000 |title=African Cichlid Fishes: Model Systems for Evolutionary Biology |journal=Annual Review of Ecology and Systematics |volume=31 |pages=163–196 |doi=10.1146/annurev.ecolsys.31.1.163}}</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|Lukuga]] River (Lake Tanganyika's outflow), but any further distribution into the [[Congo Basin|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 [[Alkali|alkaline]] and of a higher [[pH]] (which the cichlids prefer), containing a higher [[calcium]] and mineral content than the Congo's [[Acid|acidic]], [[sediment]]-rich waters, which collect much organic detritus from the surrounding [[Rainforest|rainforests]]. In areas of the Congo away from rapids or [[whitewater]], the accumulated sediment and organic matter creates sections of "[[Blackwater river|blackwater]]", with a high concentration of [[Tannin|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 [[Lake Malawi|Malawi]] or [[Lake Victoria|Victoria]]—which both have experienced relatively recent explosive species [[Adaptive radiation|radiations]] (resulting in many closely related species)—,<ref name="Turner2001">{{cite journal |last1=Turner |first1=Seehausen |last2=Knight |first2=Allender |last3=Robinson |year=2001 |title=How many species of cichlid fishes are there in African lakes? |doi=10.1046/j.1365-294x.2001.01200.x |journal=Molecular Ecology |volume=10 |issue=3 |pages=793–806 |pmid=11298988 |bibcode=2001MolEc..10..793T |s2cid=12925712 |doi-access=free}}</ref> its cichlids are the most [[Morphology (biology)|morphologically]] and [[Genetics|genetically]] diverse.<ref name=":0">{{Cite journal |last1=Meyer |last2=Matchiner |last3=Salburger |first1=Britta |first2=Michael |first3=Walter |date=25 November 2013 |title=A tribal level phylogeny of Lake Tanganyika cichlid fishes based on a genomic multi-marker approach |journal=Molecular Phylogenetics and Evolution |doi=10.1016/j.ympev.2014.10.009 |pmid=25433288 |volume=83 |pages=56–71 |pmc=4334724}}</ref><ref>{{Cite journal |last=Seehausen |first=O. |year=2015 |title=Process and pattern in cichlid radiations – inferences for understanding unusually high rates of evolutionary diversification |journal=New Phytologist |doi=10.1111/nph.13450 |volume=207 |issue=2 |pages=304–312 |pmid=25983053 |doi-access=free}}</ref> This is linked to the maturity of Tanganyika, as it is far older than the other lakes.<ref>{{cite journal |last1=Nishida |first1=M |year=1991 |title=Lake Tanganyika as an evolutionary reservoir of old lineages of East African cichlid fishes: Inferences from allozyme data |journal=Experientia |volume=47 |issue=9 |pages=974–979 |doi=10.1007/bf01929896 |s2cid=37599331}}</ref> Tanganyika has the largest number of [[Endemism|endemic]] cichlid [[genera]] of all African lakes.<ref name=":0"/> All Tanganyika cichlids are in the subfamily [[Pseudocrenilabrinae]]. Of the 10 [[Tribe (biology)|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>{{cite journal |last1=Sparks |last2=Smith |year=2004 |title=Phylogeny and biogeography of cichlid fishes (Teleostei: Perciformes: Cichlidae) |journal=Cladistics |volume=20 |issue=6 |pages=501–517 |doi=10.1111/j.1096-0031.2004.00038.x |pmid=34892958 |citeseerx=10.1.1.595.2118 |s2cid=36086310}}</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 {{cvt|100|m|ft|-1}}, but some deep-water species regularly descend to {{cvt|200|m|ft|-1}}.<ref name=Kirchberger2012>{{cite journal |last1=Kirchberger |last2=Sefc |last3=Sturmbauer |last4=Koblmuller |year=2012 |title=Evolutionary History of Lake Tanganyika's Predatory Deepwater Cichlids |journal=International Journal of Evolutionary Biology |volume=2012 |pages=716209 |doi=10.1155/2012/716209 |pmid=22675652 |pmc=3362839 |doi-access=free}}</ref> ''[[Trematocara]]'' species have, exceptionally, been found at more than {{cvt|300|m|ft|-1}}, which is deeper than any other cichlid known.<ref>Loiselle, Paul (1994). The Cichlid Aquarium, p. 304. Tetra Press, Germany. {{ISBN|978-1564651464}}.</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.&nbsp;213–214. Springer. {{ISBN|978-0792360179}}</ref> No Tanganyika cichlids are truly [[pelagic]] ''and'' offshore, except for some of the [[piscivorous]] ''Bathybates''.<ref name=Kirchberger2012/> Two of these, ''[[Bathybates fasciatus|B. fasciatus]]'' and ''[[Bathybates leo|B. leo]]'', mainly feed on [[Tanganyika sardine]]s.<ref name=Kirchberger2012/><ref name=McConnell1987/> Tanganyika cichlids differ extensively in [[ecology]], and include species that are [[herbivore]]s, [[detritivore]]s, [[planktivore]]s, [[insectivore]]s, [[molluscivore]]s, [[scavenger]]s, [[scale-eater]]s and piscivores.<ref name=MortiffTangDiv/> These dietary specializations, however, have been shown to be variable and subject to [[Opportunism|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>{{Cite journal |vauthors=Golcher-Benavides J, Wagner CE |title=Playing out Liem's Paradox: Opportunistic Piscivory across Lake Tanganyikan Cichlids |journal=The American Naturalist |year=2019 |volume=194 |issue=2 |pages=260–267 |doi=10.1086/704169 |pmid=31318283 |s2cid=155438716}}</ref> The fishes' breeding behavior falls into two main groups: the substrate- or sand-spawners (often in caves or rock crevices) and the [[mouthbrooder]]s.<ref name=Schliewen1992>Schliewen, U. (1992). Aquarium Fish. Barron's Educational Series. {{ISBN|978-0812013504}}.</ref> Among the endemic species are two of the world's smallest cichlids, ''[[Neolamprologus multifasciatus]]'' and ''[[Neolamprologus similis|N. similis]]'' (both [[shell dweller]]s), measuring up to {{cvt|4-5|cm|in}},<ref>{{FishBase | genus = Neolamprologus | species = multifasciatus | month = March| year = 2017}}</ref><ref>{{FishBase | genus = Neolamprologus | species = similis | month = March| year = 2017}}</ref> and one of the largest, the [[giant cichlid]] (''Boulengerochromis microlepis'') at up to {{cvt|90|cm|ft}}.<ref name=MortiffTangDiv/><ref name=PracFishKeepLarge>{{cite web |url=http://www.practicalfishkeeping.co.uk/features/articles/the-10-biggest-cichlids |title=The 10 biggest cichlids |publisher=Practical Fishkeeping |date=13 June 2016 |access-date=17 March 2017 |archive-date=20 March 2017 |archive-url=https://web.archive.org/web/20170320052647/http://www.practicalfishkeeping.co.uk/features/articles/the-10-biggest-cichlids |url-status=live }}</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>{{cite web |url=http://www.aquariumslife.com/featured/lake-tanganyika-biotope/ |title=tanganyika biotope aquarium |date=2010-02-10 |publisher=Aquariums Life |archive-url=https://web.archive.org/web/20120302102920/http://www.aquariumslife.com/featured/lake-tanganyika-biotope/ |archive-date=2012-03-02 |url-status=usurped |access-date=2014-02-03}}</ref>

<gallery mode="packed" heights="100" perrow="5" caption="Cichlid tribes in Lake Tanganyika (E = tribe endemic or near-endemic)">
File:Bathybates ferox.jpg|[[Bathybatini]] (E): ''[[Bathybates ferox]]'' is [[benthic]] and [[piscivorous]], but the genus also includes pelagic species.<ref name=Kirchberger2012/> The tribe is sometimes split in three, others being [[Hemibatini]] and [[Trematocarini]]<ref>{{Cite journal |last1=Meyer |last2=Matchiner |last3=Salburger |year=2015 |title=Lake Tanganyika—A 'Melting Pot' of Ancient and Young Cichlid Lineages (Teleostei: Cichlidae)? |journal=PLOS ONE |volume=10 |issue=7 |doi=10.1371/journal.pone.0125043 |pmid=25928886 |page=e0125043 |pmc=4415804 |bibcode=2015PLoSO..1025043W |doi-access=free}}</ref><ref name=Weiss2015>{{Cite journal |last1=Weiss |last2=Cotterill |last3=Schliewen |year=2015 |title=A tribal level phylogeny of Lake Tanganyika cichlid fishes based on a genomic multi-marker approach |journal=Molecular Phylogenetics and Evolution |volume=83 |pages=56–71 |doi=10.1016/j.ympev.2014.10.009 |pmid=25433288 |pmc=4334724}}</ref>
File:Benthochromis tricoti.jpg|[[Benthochromini]] (E): ''[[Benthochromis horii]]'' was scientifically described in 2008, but has often been misidentifed as ''[[Benthochromis tricoti|B. tricoti]]''<ref>{{cite journal |title=Description of a new cichlid fish species of the genus Benthochromis (Perciformes: Cichlidae) from Lake Tanganyika |last1=Takahashi |journal=Journal of Fish Biology |volume=72 |issue=3 |pages=603–613 |year=2008 |doi=10.1111/j.1095-8649.2007.01727.x |first1=T.|bibcode=2008JFBio..72..603T }}</ref>
File:DKoehl Boulengerochromis microlepis.jpg|[[Boulengerochromini]] (E): ''[[Boulengerochromis microlepis]]'' is one of the world's largest cichlids<ref name=PracFishKeepLarge/> and only member of its tribe<ref name=Weiss2015/>
File:Cyphotilapia frontosa2.jpg|[[Cyphotilapiini]] (E): ''[[Cyphotilapia frontosa]]'', one of only two similar species in the [[Tribe (biology)|tribe]]<ref>{{cite journal |last1=Takahashi |first1=T. |last2=Nakaya |first2=K. |year=2003 |title=New species of Cyphotilapia (Perciformes: Cichlidae) from Lake Tanganyika, Africa |journal=Copeia |volume=2003 |issue=4 |pages=824–832 |doi=10.1643/ia03-148.1 |s2cid=83854866}}</ref>
File:Kleinschuppiger Kaerpflingsbuntbarsch Cyprichromis microlepidotus Tierpark Hellabrunn-1.jpg|[[Cyprichromini]] (E): ''[[Cyprichromis microlepidotus]]'' and other members of this tribe are open-water [[planktivore]]s<ref>{{cite iucn |author=Bigirimana, C. |date=2006 |title=''Cyprichromis microlepidotus'' |volume=2006 |page=e.T60487A12363286 |doi=10.2305/IUCN.UK.2006.RLTS.T60487A12363286.en |access-date=13 November 2021}}</ref><ref name=Smith1998>Smith, M.P. (1998). Lake Tanganyikan Cichlids, pp. 9–10. {{ISBN|0-7641-0615-5}}</ref>
File:Ophthalmotilapia nasuta Kipili.jpg|[[Ectodini]] (E): ''[[Ophthalmotilapia nasuta]]'' (male) is [[sexually dimorphic]], males being more colorful with longer fins and nose<ref>{{Cite web |title=Ophthalmotilapia nasuta |website=Seriously Fish |url=https://www.seriouslyfish.com/species/ophthalmotilapia-nasuta/ |access-date=2023-03-11 |archive-date=11 March 2023 |archive-url=https://web.archive.org/web/20230311162708/https://www.seriouslyfish.com/species/ophthalmotilapia-nasuta/ |url-status=live }}</ref>
File:Eretmodus-sp-kavala1.jpg|[[Eretmodini]] (E): ''[[Eretmodus cyanostictus]]'' lives near the bottom in the turbulent, coastal [[surf zone]],<ref>{{FishBase | genus = Eretmodus | species = cyanostictus | month = April| year = 2017}}</ref> like other members of its tribe<ref name=Smith1998/>
File:Astatotilapia burtoni.png|[[Haplochromini]]: ''[[Astatotilapia burtoni]]'' is one of the few Tanganyika species,<ref name=FishbaseList/> unlike other [[African Great Lakes]] where most belong to this tribe<ref name=Lowe2009>{{cite journal |last1=Lowe-McConnell |first1=R |year=2009 |title=Fisheries and cichlid evolution in the African Great Lakes: progress and problems |journal=Freshwater Reviews |volume=2 |issue=2 |pages=131–151 |doi=10.1608/frj-2.2.2 |s2cid=54011001}}</ref>
File:Schachbrett-Schlankcichlide.jpg|[[Lamprologini]] (E): ''[[Julidochromis marlieri]]'' is popular in the [[aquarium]] trade where members of the genus are known as "Julies"<ref>{{Cite web |title=Julidochromis marlieri (Marlier's Julie) |website=Seriously Fish |url=https://www.seriouslyfish.com/species/julidochromis-marlieri/ |access-date=2023-03-11 |archive-date=11 March 2023 |archive-url=https://web.archive.org/web/20230311162711/https://www.seriouslyfish.com/species/julidochromis-marlieri/ |url-status=live }}</ref>
File:Gnathochromis premaxillaris.jpg|[[Limnochromini]] (E): ''[[Gnathochromis permaxillaris]]'' is a [[Planktivore|zooplanktivore]] with an unusual protractile mouth<ref>{{cite iucn |author=Bigirimana, C. |date=2006 |title=''Gnathochromis permaxillaris'' |volume=2006 |page=e.T60493A12364587 |doi=10.2305/IUCN.UK.2006.RLTS.T60493A12364587.en |access-date=13 November 2021}}</ref>
File:Perissodus microlepis juvenile in aquarium.jpg|[[Perissodini]] (E): ''[[Perissodus microlepis]]'', a specialized [[scale-eating]] species<ref>{{cite journal |last1=Stewart |first1=T.A. |last2=Albertson |first2=R.C. |year=2010 |title=Evolution of a unique predatory feeding apparatus: functional anatomy, development and a genetic locus for jaw laterality in Lake Tanganyika scale-eating cichlids |journal=BMC Biology |volume=8 |issue=1 |page=8 |doi=10.1186/1741-7007-8-8 |pmid=20102595 |pmc=2828976 |doi-access=free}}</ref>
File:Oreochromis tanganicae (Günther).jpg|[[Tilapiini]]: ''[[Oreochromis tanganicae]]'' is one of the most common coastal species found in local [[fish market]]s<ref>{{cite iucn |author=Ntakimazi, G. |date=2006 |title=''Oreochromis tanganicae'' |volume=2006 |page=e.T60625A12387918 |doi=10.2305/IUCN.UK.2006.RLTS.T60625A12387918.en |access-date=13 November 2021}}</ref>
File:TropheusspRed200.jpg|[[Tropheini]] (E): ''[[Tropheus moorii]]'' ("red" Chimba [[Polymorphism (biology)|morph]]) is highly variable and the [[Taxonomy (biology)|taxonomy]] of some of the morphs is questionable<ref>Begon, M.; and A.H. Fitter (1995). Advances in Ecological Research, vol. 26, p. 203. {{ISBN|0-12-013926-X}}</ref><ref>{{cite journal |last1=Salzburger |last2=Niederstätter |last3=Brandstätter |last4=Berger |last5=Parson |last6=Snoeks |last7=Sturmbauer |year=2006 |title=Colour-assortative mating among populations of Tropheus moorii, a cichlid fish from Lake Tanganyika, East Africa |journal=Proc Biol Sci |volume=273 |issue=1584 |pages=257–266 |doi=10.1098/rspb.2005.3321 |pmid=16543167 |pmc=1560039}}</ref><ref>{{Cite web |title=Tropheus Genus Evolution |url=http://www.cichlidworld.eu/clanky/evolucia_tropheus_e.htm |access-date=2023-03-11 |website=Cichlid World |author1=Robert Toman |date=2017 |archive-date=19 September 2021 |archive-url=https://web.archive.org/web/20210919232102/http://www.cichlidworld.eu/clanky/evolucia_tropheus_e.htm |url-status=dead }}</ref>
</gallery>