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===Fish=== ====Fishing==== [[File:Pesce ad essiccare sulla riva del lago malawi.JPG|thumb|left|[[Lake Malawi sardine]]s (''Engraulicypris sardella'') spread out to dry on the shore of the lake]] Lake Malawi has for millennia provided a major food source to the residents of its shores since its waters are rich in fish. Among the most popular are the four species of chambo, consisting of any one of four species in the subgenus ''Nyasalapia'' (''[[Oreochromis karongae]]'', ''[[Oreochromis lidole|O. lidole]]'', ''[[Oreochromis saka|O. saka]]'' and ''[[Oreochromis squamipinnis|O. squamipinnis]]''), as well as the closely related ''[[Oreochromis shiranus|O. shiranus]]''.<ref name=Turner1992>Turner, G.F.; and N.C. Mwanyama (July 1992).[http://www.fao.org/docrep/006/ad202e/AD202E00.htm Distribution and Biology of Chambo (Oreochromis spp.) in Lakes Malawi and Malombe.] [[Food and Agriculture Organization]], Fisheries and Aquaculture Department, FI:DP/MLW/86/013, Field Document 21. Retrieved 13 April 2017.</ref> Other species that support important fisheries include the [[Lake Malawi sardine]] (''Engraulicypris sardella'') and the large [[kampango]] catfish (''Bagrus meridionalis'').<ref name=Konings/> Most fishing provides food for the [[Population growth|increasing human population]] near the lake, but some are exported from Malawi. The wild population of fish is increasingly threatened by [[overfishing]] and [[water pollution]].<ref>{{Cite web|title = Preserving the Future for Lake Malawi|url = http://web.mit.edu/africantech/www/articles/Lake_Malawi.html|website = web.mit.edu|access-date = 2015-09-14}}</ref><ref name=Banda2013>{{cite news|author=Banda, M. | title = Rapid drop in Lake Malawi's water levels drives down fish stocks | url = https://www.theguardian.com/global-development/2013/may/22/lake-malawi-water-levels-fish-stocks|newspaper=The Guardian | date = 22 May 2013 | access-date = 11 April 2017}}</ref> A drop in the lake's water level represents another threat, and is believed to be driven by water extraction by the increasing human population, [[climate change]] and [[deforestation]].<ref name=Banda2013/> The chambo and kampango have been particularly overfished (the kampango declined by about 90% from 2006 to 2016,<ref>{{Cite iucn | author1 = Phiri, T.B. | author2 = Gobo, E. | author3 = Tweddle, D. | author4 = Kanyerere, G.Z. | title = ''Bagrus meridionalis'' | amends= 2018 | page = e.T60856A155041757 | year= 2019 | doi = 10.2305/IUCN.UK.2018-2.RLTS.T60856A155041757.en}}</ref> ''O. karongae'' and ''O. squamipinnis'' by about 94%, and ''O. lidole'' might already be extinct<ref>{{Cite iucn | author1 = Kanyerere, G.Z. | author2 = Phiri, T.B. | author3 = Shechonge, A. | title = ''Oreochromis karongae'' | errata= 2019 | page = e.T61293A148647939 | year= 2018 | doi = 10.2305/IUCN.UK.2018-2.RLTS.T61293A148647939.en }}</ref><ref>{{Cite iucn | author1 = Phiri, T.B. | author2 = Kanyerere, G.Z. | title = ''Oreochromis squamipinnis'' | errata= 2019 | page = e.T60760A148648312 | year= 2018 | doi = 10.2305/IUCN.UK.2018-2.RLTS.T60760A148648312.en}}</ref>) and they are now seriously [[Threatened species|threatened]].<ref>{{Cite news|author1=McKenzie, D. |author2=B. Swails | title = Rangers struggle to save endangered fish in Lake Malawi | url = http://edition.cnn.com/2010/WORLD/africa/10/27/malawi.overfishing/|publisher=CNN | date = 22 May 2013 | access-date = 11 April 2017}}</ref> The [[IUCN]] recognises 117 species of Malawi cichlids as threatened; some of these have tiny ranges and may be restricted to rocky coastlines only a few hundred metres long.<ref name=IUCNef>{{cite web|url=http://www.iucnredlist.org/initiatives/freshwater/eastafrica/geographicpatternsea |title=Geographic Patterns|url-status=dead|archive-url=https://web.archive.org/web/20181002175558/http://www.iucnredlist.org/initiatives/freshwater/eastafrica/geographicpatternsea|archive-date=2 October 2018 |access-date=9 January 2024}}</ref> {{multiple image | direction = vertical | header = Malawi cichlids | image1 = Diplotaxodon.jpg | image2 = Adult male livingstonii.png | image3 = Copadichromis azureus.jpg | image4 = Frankfurt Zoo - Aulonocara stuartgranti Usisya.jpg | image5 = Fossorochromis rostratus - mâle en aquarium 01.jpg | image6 = Pseudotropheus saulosi.jpg | image7 = Oreochromis squamipinnis.jpg | caption7 = Top to bottom:<br> 1. ''[[Diplotaxodon]]'', one of the very few cichlid genera that occurs offshore in relatively deep water.<ref name=Lowe2003/><br> 2. ''[[Nimbochromis livingstonii]]'' is a [[piscivorous]] hap that is famous for [[Apparent death|playing dead]] to lure prey close.<ref name=Konings/><ref name=Schliewen1992>Schliewen, U. (1992). Aquarium Fish. Barron's Educational Series. {{ISBN|978-0812013504}}.</ref><ref name=Elieson/><br> 3. As typical of [[utaka]], ''[[Copadichromis azureus]]'' has bright blue males (shown) and duller females that are silvery with dark spots.<ref>Elieson, M: [http://www.cichlid-forum.com/articles/c_azureus.php Copadichromis azureus.] CichlidForum. Retrieved 16 April 2017.</ref><br> 4. ''[[Aulonocara stuartgranti]]'' is part of a group of relatively peaceful species popularly known as [[peacock cichlid]]s.<ref>Elieson, M: [http://www.cichlid-forum.com/articles/peacocks.php The Peacocks of Lake Malawi.] CichlidForum. Retrieved 16 April 2017.</ref><br> 5. ''[[Fossorochromis rostratus]]'' is an "aberrant" hap that often sifts mouthfuls of sand to extract small food organisms.<ref name=Konings/><ref>O'Brien, R: [http://www.cichlid-forum.com/articles/f_rostratus.php Fossorochromis rostratus.] CichlidForum. Retrieved 16 April 2017.</ref><br> 6. Like many [[mbuna]], ''[[Pseudotropheus saulosi]]'' is a small cichlid where both male (blue and black) and female (yellow) are colorful.<ref>Barber, P: [http://www.cichlid-forum.com/articles/p_saulosi.php Pseudotropheus saulosi.] CichlidForum. Retrieved 16 April 2017.</ref><br> 7. ''[[Oreochromis squamipinnis]]'' is one of only six native [[tilapia]] species in the lake, but these are important to fisheries. Notice the white genital tassels, longer when fully extended and unique to male chambo.<ref name=Konings/><ref name=Turner1992/> }} ====Cichlids==== Lake Malawi is noted for being the site of [[evolutionary radiation]]s among several groups of animals, most notably [[cichlid]] fish.<ref>{{cite journal | last=Svardal | first=Hannes | last2=Quah | first2=Fu Xiang | last3=Malinsky | first3=Milan | last4=Ngatunga | first4=Benjamin P | last5=Miska | first5=Eric A | last6=Salzburger | first6=Walter | last7=Genner | first7=Martin J | last8=Turner | first8=George F | last9=Durbin | first9=Richard | title=Ancestral Hybridization Facilitated Species Diversification in the Lake Malawi Cichlid Fish Adaptive Radiation | journal=Molecular Biology and Evolution | volume=37 | issue=4 | date=2020-04-01 | issn=0737-4038 | pmid=31821500 | pmc=7086168 | doi=10.1093/molbev/msz294 | pages=1100–1113|biorxiv=10.1101/738633|s2cid=202010546|hdl=1983/40d3dafd-f1d5-4ea4-a447-126df76a0651|hdl-access=free}}</ref> There are at least 700 cichlid species in Lake Malawi,<ref name=Turner2001/> with some estimating that the actual figure is as high as 1,000 species.<ref name=panda/><ref>Kornfield, I.; & P.F. Smith (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]].</ref> The actual number is labelled with some uncertainty because of the many [[undescribed species]] and the extreme variation among some species, making the task of delimiting them very complex.<ref name=Turner2001/><ref name=Konings/> Except for four species (''[[Astatotilapia calliptera]]'', ''[[Coptodon rendalli]]'', ''Oreochromis shiranus'' and ''[[Serranochromis]] robustus''), all cichlids in the lake are [[Endemism|endemic]] to the Malawi system, which also includes nearby smaller [[Lake Malombe]] and the upper Shire River.<ref name=Konings/><ref name=OliverTilapia>Oliver, M.K. (12 April 2015). [http://malawicichlids.com/mw10100.htm The Tilapias of Lake Malawi.] MalawiCichlids. Retrieved 13 April 2017.</ref><ref>Oliver, M.K. (12 April 2015). [http://malawicichlids.com/mw10000.htm The Nonendemic Haplochromine Cichlids of Lake Malawi.] MalawiCichlids. Retrieved 13 April 2017.</ref> Many of these have become popular among [[aquarium]] owners due to their bright colors. Recreating a Lake Malawi biotope to host cichlids became quite popular in the aquarium hobby.<ref>{{cite web|last1=Pardee|first1=Keith|title=African Cichlids, Lake Malawi|url=http://www.aquariumlife.net/profiles/african-cichlids-lake-malawi|website=www.aquariumlife.net|access-date=30 September 2017|language=en|archive-url=https://web.archive.org/web/20171012113129/http://www.aquariumlife.net/profiles/african-cichlids-lake-malawi|archive-date=12 October 2017|url-status=dead}}</ref> Most Malawi cichlids are found in relatively shallow coastal waters,<ref name=Konings/> but ''[[Diplotaxodon]]'' has been recorded down to depths of {{convert|200-220|m|ft|abbr=on}} and several (especially ''Diplotaxodon'', ''[[Rhamphochromis]]'' and ''[[Copadichromis quadrimaculatus]]'') are known from [[pelagic]] waters.<ref name=Lowe2003/> The cichlids of the lake are divided into two groups and the vast majority of the species are [[haplochromines]]. The [[sister species]] to the Malawi haplochromines is ''Astatotilapia'' sp. Ruaha (a currently undescribed species from [[Great Ruaha River]]), and these two separated between 2.13 and 6.76 million years ago (mya).<ref name=Genner2015>Genner; Ngatunga; Mzighani; Smith; and Turner (2015). Geographical ancestry of Lake Malawi’s cichlid fish diversity. Biol. Lett. 11: 2015023. {{doi|10.1098/rsbl.2015.0232}}</ref> The earliest divergence within the Malawi haplochromines occurred between 1.20 and 4.06 mya,<ref name=Genner2015/> but most radiations in this group are far younger; in extreme cases species may have diverged only a few hundred years ago.<ref name=Givnish1997/> The Malawi haplochromines are [[mouthbrooder]]s, but otherwise vary extensively in general behaviour and ecology.<ref name=Konings/> Within the Malawi haplochromines there are two main groups, the haps and the [[mbuna]]. The haps (they were formerly included in ''[[Haplochromis]]'') can be further subdivided into three subgroups: The relatively large, often more than {{convert|20|cm|in|abbr=on|0}} long, and aggressive [[piscivore]]s that roam various habitats in pursuit of prey, the open-water (although often not far from sand or rocks) [[utaka]] that feed in [[Shoaling and schooling|schools]] on [[zooplankton]] and typically are of medium size, and finally a subgroup of "aberrant" species that essentially are defined by them not fitting clearly into the other subgroups.<ref name=Konings/><ref name=Elieson>Elieson, M: [http://www.cichlid-forum.com/articles/haps_vs_mbuna.php Haps Vs. Mbuna.] Retrieved 11 April 2017.</ref><ref name=aquaticcom>Aquaticcommunity (2004–08).[http://www.aquaticcommunity.com/cichlid/haplochromis.php Haplochromis.] Retrieved 13 April 2017.</ref> Adult male haps generally display bright colors, while juveniles of both sexes and adult females typically show a silvery or grey coloration with sometimes irregular black bars or other markings.<ref name=Konings/><ref name=Elieson/> The second main haplochromine group are the mbuna, a name used both locally and popularly, which means "rockfish" in [[Tonga (Nyasa) language|Tonga]].<ref>Loiselle, P.V. (1988). A Fishkeepers Guide to African Cichlids, p. 97. Salamander Books, London & New York. {{ISBN|0-86101-407-3}}.</ref> They are found at rocky outcrops, [[Territory (animal)|territorially]] aggressive (although commonly found in high densities) and often specialised [[aufwuchs]] feeders.<ref name=Konings/><ref name=Elieson/> The mbuna species tend to be relatively small, mostly less than {{convert|13|cm|in|abbr=on|0}} long, and often both sexes are brightly colored with males having egg-shaped yellow spots on their [[anal fin]] (a feature particularly prevalent in the mbuna, but not exclusive to this group).<ref name=Konings/><ref name=Elieson/> The second group, the [[tilapia]], comprises only six species in two genera in Lake Malawi: The [[redbreast tilapia]] (''Coptodon rendalli''), a widespread African species, is the only substrate-spawning cichlid in the lake.<ref name=Konings/><ref>Oliver, M.K. (12 April 2015). [http://malawicichlids.com/mw10003.htm Coptodon rendalli.] Malawicichlids. Retrieved 13 April 2017.</ref> This large cichlid mainly feeds on [[macrophyte]]s.<ref name=Konings/><ref>{{FishBase species | genus = Coptodon | species = rendalli | month = April| year = 2017}}</ref> The remaining are five mouthbrooding species of ''[[Oreochromis]]''; four chambo in the subgenus ''Nyasalapia'' (''O. karongae'', ''O. lidole'', ''O. saka'' and ''O. squamipinnis'') that are endemic to the Lake Malawi system, as well as the closely related ''O. shiranus'', which also is found in [[Lake Chilwa]].<ref name=Konings/><ref name=Turner1992/><ref name=OliverTilapia/> The Malawi ''Oreochromis'' mainly feed on [[phytoplankton]], reach lengths up to {{convert|26-42|cm|in|abbr=on}} depending on the exact species, and are mostly black or silvery-gray with relatively indistinct dark bars.<ref name=Konings/><ref name=OliverTilapia/><ref>{{FishBase genus | genus = Oreochromis | month = April| year = 2017}}</ref> Male chambo have unique genital tassels when breeding, which aid in egg fertilisation in a manner comparable to the egg-spots on the anal fin of haplochromines.<ref name=Konings/><ref name=Turner1992/> ====Non-cichlids==== [[File:Catfish in Lake Malawi.jpg|thumb|left|The [[kampango]] (''Bagrus meridionalis''), one of the largest catfish, reaching up to {{convert|1.5|m|ft|abbr=on}} in length<ref>{{FishBase species | genus = Bagrus | species = meridionalis | month = April| year = 2017}}</ref>]] The vast majority of the fish species in the lake are cichlids. Among the non-cichlid native fish are several species of [[cyprinids]] (in genera ''[[Barbus]]'', ''[[Labeo]]'' and ''[[Opsaridium]]'', and the Lake Malawi sardine ''Engraulicypris sardella''), [[airbreathing catfish]] (''[[Bathyclarias]]'' and ''[[Clarias]]'', and the kampango ''Bagrus meridionalis''), [[Mochokidae|mochokid catfish]] (''[[Chiloglanis]]'' and [[Malawi squeaker]] ''Synodontis njassae''), ''[[Mastacembelus]]'' spiny eel, [[mormyrids]] (''[[Marcusenius]]'', ''[[Mormyrops]]'' and ''[[Petrocephalus]]''), the [[African tetra]] ''[[Brycinus imberi]]'', the [[poeciliid]] ''[[Aplocheilichthys johnstoni]]'', the [[spotted killifish]] (''Nothobranchius orthonotus''), and the [[mottled eel]] (''Anguilla nebulosa'').<ref name=Konings/> At a genus level, most of these are widespread in Africa, but ''Bathyclarias'' is entirely restricted to the lake.<ref>Anseaume, L.; and G.G. Teugels (1999). On the rehabilitation of the clariid catfish genus Bathyclarias endemic to the East African Rift Lake Malawi. Fish Biology 55(2): 405–419. [[doi: 10.1111/j.1095-8649.1999.tb00687.x]]</ref>
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