Fisheries and Cichlid Evolution in the African Great Lakes: Progress and Problems
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131 Article Fisheries and cichlid evolution in the African Great Lakes: progress and problems Rosemary Lowe-McConnell St George’s Park, Burgess Hill, Sussex RH15 0SG, UK. Email: [email protected] Received 22 May 2009; accepted 25 September 2009; published 16 November 2009 Abstract This paper updates Recent Advances in the African Great Lakes: Fisheries, Biodiversity and Cichlid Evolution published by the Freshwater Biological Association in Freshwater Forum (Lowe- McConnell, 2003). Since 2003 many international teams have continued research on lakes Malawi, Victoria and Tanganyika. This review discusses the decline of the important commercial fisheries in all three lakes, together with changes in ecological and limnological conditions which the fishes now face. It also describes advances in our understanding of how the spectacular flocks of endemic cichlid species have evolved in each lake and continue to coexist. Keywords: African lakes; cichlid evolution; cichlid ecology; fisheries; Lake Victoria; Lake Malawi; Lake Tanganyika. Introduction obtain funds for projects on lakes Malawi, Tanganyika and Victoria. These lakes were all facing serious threats The East African Great Lakes have long been renowned for caused by the rapid rise in human populations, with fisheries of vital importance for the rapidly rising riparian associated over-fishing, sedimentation, and pollution from human populations and as biodiversity hotspots with changes in land use in the lake basins. Since the early 1990s spectacular endemic faunas. The flocks of cichlid fishes research involving over a hundred scientists, financed unique to each of the three largest lakes, Victoria (69 000 by many international bodies, has produced numerous km2) and the long deep rift-valley lakes, Tanganyika and publications in widely-scattered journals. In 2003, the Malawi (previously Nyasa) (Fig. 1), have offered unique Freshwater Biological Association published a Special opportunities to investigate how new species evolve and Issue of Freshwater Forum on Recent Advances in the African coexist (as discussed by Lowe-McConnell, 1993, 1996). Great Lakes: Fisheries, Biodiversity and Cichlid Evolution In 1989, an International Symposium on ‘Resource Use (Lowe-McConnell, 2003). This synopsis summarised the and Conservation of the African Great Lakes’ was held at status of the fisheries and the then current ideas on how Bujumbura on Lake Tanganyika, followed by numerous the rich endemic fish faunas had evolved. It also described workshops at other venues. Further international interest the biodiversity surveys made in each lake based on in biodiversity, generated at the Rio Earth Summit in 1992, underwater (SCUBA) observations of fish ecology and then enabled the United Nations and other agencies to behaviour, followed by cichlid breeding experiments in DOI: 10.1608/FRJ-2.2.2 Freshwater Reviews (2009) 2, pp. 131-151 © Freshwater Biological Association 2009 132 Lowe-McConnell, R. Fig. 1. The East African Great Lakes: Lake Victoria drains via Lake Kyoga and Lake Albert to the Nile, Lake Tanganyika to the Congo system (Zaire) and Lake Malawi to the Zambezi River. Figure originally from Lowe-McConnell, R. 1996. Fish communities in the African Great Lakes. Environmental Biology of Fishes, 45: 219-235. Reproduced with kind permission of Springer Science and Business Media. aquaria and DNA techniques in many laboratories to test ideas on the evolution, origins and relationships of the Fisheries cichlid species. New publications since 2003 have further advanced our understanding of how the spectacular Lake Malawi fisheries flocks of cichlid species have evolved and coexist in lakes Victoria and Malawi, but the main commercial fisheries Sixty years ago the first fishery survey of Lake Nyasa (now of these lakes, so needed to feed the rapidly rising human L. Malawi) showed that the main commercial fisheries populations, have declined. The present review describes were supported by cichlid tilapia caught in shore seines the most outstanding contributions on both these topics and open water ring nets (Bertram et al., 1942). Since published from 2003 to 2009. then, however, tilapia abundances have declined and the © Freshwater Biological Association 2009 DOI: 10.1608/FRJ-2.2.2 Fisheries and cichlid evolution in the African Great Lakes 133 species of endemic tilapia are now so scarce that the main could adversely affect Malawi’s endemic fishes, as it did commercial fishery (Maldeco, which works in association when introduced into Madagascar (Reinthal et al., 2003). with the Malawi Fishery Department) has resorted to Research into environmental conditions affecting fish breeding tilapia in fish-farm ponds to stock large enclosures production in Lake Malawi has included, for example, fixed in open waters in the south east arm of the lake. the work by Duponchelle et al. (2005) on food partitioning Here they rear endemic tilapia (Oreochromis shiranus and within the species-rich benthic fish community. Using a ‘chambo’ O. karongae) but the endemic, more open-water- combination of stable isotopes and stomach analyses, they dwelling O. lidole, once so abundant in ring net catches, has found that, although benthic algal production contributed been over fished to such an extent that it has now vanished to the energy requirements of offshore fishes living in and is ‘probably extinct’ (Turner, personal communication). water 10 m to 30 m deep, the larvae of the abundant lake Maldeco, which produces 70 % of Malawi’s total fish catch, fly Chaoborus edulis were the most important food source no longer targets tilapia (the most popular fish) which was for demersal fishes, thus supporting the hypothesis reported in 2005 to be at the ‘lowest point ever’. Artisanal that demersal fish production in L. Malawi is sustained fisheries for cichlids and other fishes continue elsewhere on mainly through the pelagic food chain, rather than from this 600 km long lake, from which total catches are reported benthic detritus. Isotopic differences among species by the Malawi Fisheries Department to fluctuate between with apparently similar diets, feeding behaviour and 26 000 and 47 000 tonnes annually. To augment the fish depth preferences, suggested that important resource supplies Maldeco uses three bottom trawlers which catch partitioning does exist among L. Malawi’s benthic mainly very numerous small haplochromine cichlid species, haplochromine cichlids. A Special Issue of the Journal of including ndunduma Diplotaxodon of several species, Great Lakes Research on The African Great Lakes (eds Bootsma discovered during the UK/SADC Pelagic Zone survey in et al., 2003), has an introductory paper by Bootsma & 1995. Small cichlid species now contribute between 60 % Hecky (2003) comparing the biology and limnology of and 70 % of Maldeco’s annual catch, of which 90 % is sold the African Great Lakes. Twelve of the other very diverse fresh (frozen); Maldeco also has a fish processing plant and papers were based on data from L. Malawi. These, in smoke kiln for fillets of any large fish they catch, especially addition to discussing seasonal and spatial patterns of catfishBagrus meridionalis. To help meet the country’s large experimental trawl catches (Deponchelle et al., 2003) demand for fish, they have acquired another landing site at and the return to local fisheries management (Dobson & Salima on the main lake, for which a fourth fishing vessel is Lynch, 2003), included other observations by numerous on order (Press Corporation Ltd, 2009). authors on meteorology, nutrient upwelling, sources Fish from smaller lakes in Malawi, and from pond and fluxes of organic carbon, silica cycling, community culture, also contribute to the country’s total catch. The composition, distribution and nutrient status of epilithic WorldFish Centre, based in the Philippines, has assisted periphyton in the rocky littoral, environmental factors Malawian farmers to grow tilapia in rain-fed ponds for controlling the distributions of benthic invertebrates on which new varieties of tilapia are being selected for fast rocky shores, and nitrogen and phosphorus regeneration growth. Commendably, Malawi does not stock exotic fish, by cichlids in the littoral zone. In a paper on the impact which can so easily escape to the detriment of indigenous of land use on sediments and nutrients, Hecky et al. species. The very vigorous Nile tilapia (O. niloticus), (2003) showed that changes in land use in the L. Malawi cultivated in many ponds worldwide, when introduced into basin, with its rapidly increasing human population and Lake Victoria ousted the endemic tilapia (as described later). associated agricultural development and forest clearance, This aggressive species has now spread in southern Africa, probably increased nutrient loading to the lake by 50 %. including Lake Kariba from where it has replaced Zambezi Responses to nutrient-enrichment experiments tilapias (Tweddle, 2007). This very dominant tilapia species (with P, N and Fe) demonstrated that, in adequate light, DOI: 10.1608/FRJ-2.2.2 Freshwater Reviews (2009) 2, pp. 131-151 134 Lowe-McConnell, R. phytoplankton quickly become nutrient-deficient. O. niloticus thrived despite the presence of this predatory Enrichment experiments with and without zooplankton centropomid (now Latidae) that was introduced into the (> 50 µm) by Guildford et al. (2003) in Lake Malawi Ugandan waters of the lake in the mid 1950s as a ‘sport’ fish (during three seasons – stratified rainy, deep mixing and and to crop the numerous