Changes in the prey ingested and the variations in the Nile perch and other fish stocks of Lake Kyoga and the northern waters of Lake Victoria (Uganda)

The fish stocks of Lakes Kyoga and Victoria have changed since Nile perch, Lates niloticus (L.), was introduced, and this is reflected in the prey ingested by the predator. Initially, haplochromine cichlids constituted the main prey of most sizes of Nile perch. As the stocks of these have declined, Caridina nilotica (Roux) and Anisopteran nymphs have become the dominant food of the juveniles, while Rastrineobola argentea (Pellegrin), juvenile Nile perch and Oreochromis niloticus (L.) have become the main food of larger Nile perch. Apart from R. argentea , most of the native fish species of these lakes have disappeared. The stocks of Nile perch in Lake Kyoga, to which it was introduced earlier than to Lake Victoria, have declined after dominating the fishery since 1965. and have been superseded by O. niloricus . an introduced herbivore. Similar changes are now occurring in Lake Victoria. The Nile perch might not maintain the high yield realized in the two lakes when haplochromines were abundant. It is therefore necessary to exercise caution with high and long-term investments aimed specifically at developing the Nile perch fishery.     

Trophic Niche Segregation in the Nilotic Ichthyofauna of Lake Albert (Uganda, Africa)

Synopsis Nile perch, Lates niloticus, and Nile tilapia, Oreochromis niloticus, were originally transplanted from Lake Albert in western Uganda to the African Great Lakes, Lake Victoria and Lake Kyoga, where they are partially implicated in reduction of the fish species diversity. Lake Albert is facing multiple environmental changes, including declining fish species diversity, hyper-eutrophication, hypoxia, and reduced fish catches. To examine the role of Nile perch and Nile tilapia in the food web in their native Lake Albert, we estimated their diets using stable nitrogen and carbon isotopes. In Lake Albert, the tilapiine congeners (closely related species), Tilapia zillii, Oreochromis leucostictus, and Sarethorodon galilaeus, and the centropomid Nile perch congener, Lates macrophthalmus, have narrower diet breath in the presence of the native O. niloticus and L. niloticus. A computerized parameter search of dietary items for five commercially important fish species (Hydrocynus forskahlii, Bagrus bayad, L. niloticus, Alestes baremose and Brycinus nurse) was completed using a static isotopic mixing model. The outcome of the simulation for most fish species compared favorably to previously published stomach contents data for the Lake Albert fishes dating back to 1928, demonstrating agreement between stable isotope values and analyses of stomach contents. While there were some indications of changes in the diets of L. niloticus and A. baremose diets over the past 20 years in parallel with other changes in the lake, for the most part, food web structure in this lake remained stable since 1928. The Lake Albert fish assemblage provides insight into the invasion success of L. niloticus and O. niloticus.     

The tilapiine fish stock of Lake Victoria before and after the Nile perch upsurge

Since the beginning of fisheries in Lake Victoria, two native tilapiine species, Oreochromis esculentus and Oreochromis variabilis , were the main target of the local fishermen. A continuous increase in fishing pressure led initially to a declining catch per unit of effort, and a smaller average fish size; eventually, there was a reduced landing of tilapiines. To boost the fisheries, three alien tilapiine species and the Nile perch Lates niloticus were introduced. Thirty years after its introduction, Oreochromis niloticus appeared to be the most successful tilapiine species. It replaced the indigenous tilapiines almost completely before the Nile perch came to dominate the ecosystem of Lake Victoria. Reduced fishing pressure on the tilapiines in the 1980s, due to the shift of the local fishery towards the Nile perch, resulted in an increase in the stock of O. niloticus and an increase in average fish size. Subsequently, the total mass of O. niloticus landed increased. The stocks of the indigenous tilapiines did not recover but declined to extremely low levels, or vanished from the main lake. Currently, these species still occur in satellite lakes of Lake Victoria, from which O. niloticus is absent. Nile perch feed on O. niloticus; however, the limited overlap in distribution between piscivorous Nile perch and O. niloticus of consumable sizes is probably an important factor in explaining the coexistence of the two species. The main cause of the disappearance of the native tilapiine species is presumed to be competitive dominance by O. niloticus .     

Co-introduction of Dolicirroplectanum lacustre,a monogenean gill parasite of the invasive Nile perch Lates niloticus: intraspecific diversification and mitonuclear discordance in native versus introduced areas

The Nile perch (Lates niloticus) is a notorious invasive species. The introductions of Nile perch into several lakes and rivers in the Lake Victoria region led to the impoverishment of trophic food webs, particularly well documented in Lake Victoria. Additionally, its parasites were co-introduced, including Dolicirroplectanum lacustre (Monogenea, Diplectanidae). Dolicirroplectanum lacustre is the single monogenean gill parasite of latid fishes (Lates spp.) inhabiting several major African freshwater systems. We examined the intra-specific diversification of D. lacustre from Lates niloticus in Lake Albert, Uganda (native range) and Lake Victoria (introduced range) by assessing morphological and genetic differentiation, and microhabitat preference. We expected reduced morphological and genetic diversity for D. lacustre in Lake Victoria compared with Lake Albert, as a result of the historical introductions. We found that D. lacustre displayed high morphological variability within and between African freshwaters, with two morphotypes identified, as in former studies. The single shared morphotype between Lake Albert and Lake Victoria displayed similar levels of haplotype and nucleotide diversity between the lakes. Mitonuclear discordance within the morphotypes of D. lacustre indicates an incomplete reproductive barrier between the morphotypes. The diversification in the mitochondrial gene portion is directly linked with the morphotypes, while the nuclear gene portions indicate conspecificity. Based on our results, we reported reduced genetic and morphological diversity, potentially being a result of a founder effect in Lake Victoria.     

Genetic structure and gene flow in an endangered native tilapia fish (Oreochromis esculentus) compared to invasive Nile tilapia (Oreochromis niloticus) in Yala swamp, East Africa

The introduction of invasive Nile tilapia (Oreochromis niloticus), and the rapacious predator Nile perch (Lates niloticus), into Lake Victoria resulted in a decline in population sizes, genetic diversity and even extirpation of native species which were previously the mainstay of local fisheries. However, remnant populations of native fish species, including tilapia, still persist in satellite lakes around Lake Victoria where they may coexist with O. niloticus. In this study we assessed population genetic structure, diversity, and integrity of the native critically endangered Singidia tilapia (O. esculentus) in its refugial populations in the Yala swamp, Kenya, and contrasted this diversity with populations of the invasive tilapia O. niloticus in satellite lakes (Kanyaboli, Namboyo and Sare) and Lake Victoria. Based on mtDNA control region sequences and eight nuclear microsatellite loci, we did not detect any mtDNA introgression between the native and the invasive species in Lakes Kanyaboli and Namboyo, but did find low levels of nuclear admixture, primarily from O. niloticus to O. esculentus. Some genetic signal of O. esculentus in O. niloticus was found in Lake Sare, where O. esculentus is not found, suggesting it has recently been extirpated by the O. niloticus invasion. In both species, populations in the satellite lakes are significantly genetically isolated from each other, with private mtDNA haplotypes and microsatellite alleles. For O. niloticus, genetic diversity in satellite lakes was similar to that found in Lake Victoria. Our data imply a low frequency of immigration exchange between the two populations of O. esculentus and we suggest that the populations of this endangered species and important fisheries resource should be conserved separately in Lakes Kanyaboli and Namboyo and with high priority.     

Trophic shifts in predatory catfishes following the introduction of Nile perch into Lake Victoria

This study looked for evidence of trophic shifts in the diet of two predatory catfishes ( Bagrus docmac and Schilbe intermedius ) following the establishment of introduced Nile perch ( Lates niloticus ) into lakes of the Lake Victoria basin. Bagrus docmac exhibited a shift from a primarily piscivorous diet dominated by haplochromine cichlids to a broader diet that included a significant proportion of invertebrates and the cyprinid fish, Rastrineobola argentea , which became abundant following depletion of the haplochromines. Schilbe intermedius exhibited a trophic shift from a piscivorous diet dominated by haplochromines to an insectivorous diet. The flexibility in diet exhibited by these two catfishes may have permitted these species to persist, albeit in reduced numbers, subsequent to the introduction of Nile perch and may facilitate resurgence as fishing pressure reduces numbers of large Nile perch.     

The food of the Nile perch, Lates niloticus (L.), after the disappearance of the haplochromine cichlids in the Nyanza Gulf of Lake Victoria (Kenya)

After the disappearance of the haplochromine species in the Nyanza Gulf of Lake Victoria as a result of predation by Lates niloticus , the latter has turned its attention to aquatic invertebrates and other fish. Changes in the diet of the Nile perch with increase in its size have been observed: young L. niloticus preyed mostly on invertebrates, including crustaceans and various small aquatic insects; large, immature L. niloticus supplemented the invertebrate diet with both young and small fish; adults above 80 cm total length were mainly piscivorous. L. niloticus feeds on fish prey of about one third its own length.
The tendency of L. niloticus to switch from one prey item to another, depending on availability, is reported; e.g., in the Nyanza Gulf, the prey diet has shifted from the haplochromine to Caridina nilotica and L. niloticus juveniles.     

The decline of the native fishes of lakes Victoria and Kyoga (East Africa) and the impact of introduced species,especially the Nile perch,Lates niloticus,and the Nile tilapia,Oreochromis niloticus

Synopsis There has been a decline, and in some cases an almost total disappearance, of many of the native fish species of lakes Victoria and Kyoga in East Africa since the development of the fisheries of these lakes was initiated at the beginning of this century. The Nile perch, Lates niloticus, a large, voracious predator which was introduced into these lakes about the middle of the century along with several tilapiine species, is thought to have caused the reduction in the stocks of several species. But overfishing and competition between different species also appear to have contributed to this decline. By the time the Nile perch had become well established, stocks of the native tilapiine species had already been reduced by overfishing. The Labeo victorianus fishery had also deteriorated following intensive gillnetting of gravid individuals on breeding migrations. L. niloticus is, however, capable of preying on the species which haven been overfished and could have prevented their stocks from recovering from overfishing. L. niloticus is also directly responsible for the decline in populations of haplochromine cichlids which were abundant in these lakes before the Nile perch became established. Even without predation by Nile perch, it has been shown that the haplochromine cichlids could not have withstood heavy commercial exploitation if a trawl fishery had been established throughout Lake Victoria. Their utilisation for human food has also posed some problems. The abundance of the native tilapiine species may also have been reduced through competition with introduced species which have similar ecological requirements. At present, the Nile perch and one of the introduced tilapiine species, Oreochromis niloticus, form the basis of the fisheries of lakes Victoria and Kyoga.Invited editorial     

The use of integrated constructed wetlands (ICW) for the treatment of separated swine wastewaters

Haplochromine cichlids used to be the main prey of the introduced Nile perch, Lates niloticus, in Lake Victoria. After depletion of the haplochromine stocks at the end of the 1980s, Nile perch shifted to the shrimp Caridina nilotica and to a lesser degree to its own young and the cyprinid Rastrineobola argentea. In the present study, we investigated the Nile perch diet in the northern Mwanza Gulf after resurgence of some of the haplochromine species and compared it with data collected in the same area in 1988/1989. It became clear that haplochromines are again the major prey of Nile perch. The dietary shift from invertebrate feeding (shrimps) to feeding on fish (haplochromine cichlids) occurs at a smaller size than it did when Nile perch were taking primarily dagaa and juvenile Nile perch as their fish prey. The apparent preference for haplochromines as prey has reduced the degree of cannibalism considerably, which may have a positive impact on Nile perch recruitment.     

The impact of the introduction of Nile perch, Lates niloticus (L.) on the fisheries of Lake Victoria

The piscivorous Nile perch was introduced into Lake Victoria some 30 years ago, since when it has completely transformed the fishing industry and the species composition of the fish fauna of the lake. The original multispecies fishery, based mostly on cichlids (haplochromines, tilapias), cyprinids ( Barbus, Labeo, Rastrineobola ) and siluroids ( Bagrus, Clarias, Synodontis, Schilbe ), has changed dramatically to one based on three species: the introduced Nile perch, the cyprinids, Rastrineobola argenrea (Pellegrin), and the introduced Nile tilapia, Oreochromis niloticus (Linnaeus).
Within 25 years of its introduction the Nile perch became ubiquitous and now occurs in virtually every habitat with the exception of swamps and affluent rivers. It has preyed on all other species with profound effects, especially on the stocks of haplochromines. These originally comprised 80% of the total fish biomass in Lake Victoria, but have now decreased to less than 1% offish catches from the Kenyan waters of the lake. The fishermen of Lake Victoria have adjusted to this ecological crisis by using large-meshed nets to catch Nile perch, which has become the most important commercial species. For the first time in the history of Lake Victoria, fish fillets are now being exported to several overseas countries: the fillets are all from Nile perch.     

Management of the Nile perch, Lates niloticus fishery in Lake Victoria in light of the changes in its life history characteristics

Nile perch was introduced into lakes Victoria and Kyoga in the 1950s and 1960s from Lake Albert. The changes in prey eaten and the life history characteristics of Nile perch in lakes Victoria and Kyoga from the 1960s to 1990s were examined and compared with Lake Albert. The dominant prey eaten changed from haplochromines, Caridina nilotica, Rastrineobola argentea, and Nile perch juveniles. The condition factor deteriorated from 1.4 in 1960s to 1.2 in 1990s, compared with 1.3 in Lake Albert suggesting a reduction in food supply. Therefore, exploitation of Nile perch prey should be controlled. The size at first maturity increased from 30–40 to 40–50 cm and 50–59 to 80–100 cm for males and females, which is similar to Lake Albert. Sex ratios decreased from 85–100 to 20–65 females for every 100 males suggesting that Nile perch had less capacity to replenish its stocks such that breeding females should be protected. As males mature at 50–55 cm and females at 80–100 cm, immature males of <50 cm and breeding females of >100 cm could be protected through size selective exploitation of fish of 50–100 cm using gillnets of 127–254 mm.     

尼罗尖吻鲈和鳜鱼染色体组型分析及比较

采用PHA、秋水仙碱腹腔或背部肌肉注射,活体培养法,以前肾为材料,低渗-空气干燥法制片,进行染色体观察,运用Micromeasure version 3.3染色体分析软件和Photoshop 7.0软件首次分析了尼罗尖吻鲈的染色体数目和核型,并同鳜鱼染色体数目和核型进行了分析比较,对今后拟采取的杂交尝试提供理论基础。结果显示：尼罗尖吻鲈染色体众数为2n=48,核型公式为2m+4sm+12st+30t,染色体臂数（NF）为54;鳜鱼染色体众数为2n=48,核型公式为6sm+12st+30t,染色体臂数（NF）亦为54;两种鱼染色体短臂上均无随体,单臂染色体较多。分析表明尼罗尖吻鲈与鳜鱼杂交成功的可能性较大。     

Divergence in aerobic scope and thermal tolerance is related to local thermal regime in two populations of introduced Nile perch (Lates niloticus)

We tested whether thermal tolerance and aerobic performance differed between two populations of Nile perch (Lates niloticus) originating from the same source population six decades after their introduction into two lakes in the Lake Victoria basin in East Africa. We used short-term acclimation of juvenile fish to a range of temperatures from ambient to +6°C, and performed critical thermal maximum (CT_max) and respirometry tests to measure upper thermal tolerance, resting and maximum metabolic rates, and aerobic scope (AS). Across acclimation temperatures, Nile perch from the cooler lake (Lake Nabugabo, Uganda) tended to have lower thermal tolerance (i.e., CT_max) and lower aerobic performance (i.e., AS) than Nile perch from the warmer waters of Lake Victoria (Bugonga region, Uganda). Effects of temperature acclimation were more pronounced in the Lake Victoria population, with the Lake Nabugabo fish showing less thermal plasticity in most metabolic traits. Our results suggest phenotypic divergence in thermal tolerance between these two introduced populations in a direction consistent with an adaptive response to local thermal regimes.     

Effects of Nile perch, Lates niloticus, on functional and specific fish diversity in Uganda's Lake Kyoga system

The introduction of Nile perch, Lates niloticus, to Lake Victoria, East Africa, interacted with eutrophication to cause a reorganization of the lake's food web and the extirpation of many endemic fishes. The Lake Kyoga satellite system lies downstream from Lake Victoria. It encompasses species‐rich lakes where Nile perch are absent or very rare, and low diversity lakes where L. niloticus is abundant. In 1999 we surveyed seven lakes in the Kyoga system using experimental monofilament gill nets (1/4–1 inches variable mesh). At Boston University we assessed δ¹⁵N signatures of epaxial muscle from subsamples of the catch (n = 361). These signatures are often highly correlated with the near‐term mean realized trophic position of an individual organism. A neural network analysis of fish length, species name, trophic level, and lake of origin fish explained 94% of the sample variance in δ¹⁵N. We analysed statistical patterns in these signatures at a number of spatial scales. The relationship between trophic level and δ¹⁵N varied greatly among lakes. Higher diversity perch‐free lakes had greater variance in δ¹⁵N values and fish lengths than lower diversity Nile perch lakes, suggesting an important relationship between species diversity and functional diversity. Against expectations, lake size was negatively correlated with δ¹⁵N. Patterns in stable isotope signatures indicated that Nile perch lakes have shorter food chains than perch‐free lakes. The results throw up two management problems for the Kyoga system. Impacted lakes need to be studied to understand and ameliorate the community‐level effects of Nile perch introduction, whereas the species‐rich nonperch lakes, which harbour a large proportion of the remaining diversity of regionally endemic taxa, are in need of conservation planning.     

Human-induced changes in the composition of fish communities in the African Great Lakes

Summary Large natural African lakes contain unique and diverse fish faunas which have evolved within each lake in a comparatively short period of time. members of the family Cichlidae are particularly diverse, although there is strong evidence to show that the haplochromines in Lake Victoria, and possibly Lake Malawi, are monophyletic. The unique faunas in Lakes Victoria and Kyoga have been subject to perturbations from the introduction of exotic fish, and the faunas in these and other lakes have been disturbed by fishing activities and other human endeavours.Factors governing the establishment of exotic species are not clearly understood. The exotic fish must be physiologically adapted to their new environment, able to compete successfully both for habitat and for food at each stage of their life history, able to avoid predation and must have a suitable reproductive potential. Although about 50 species of fish have been introduced into African inland waters, including reservoirs, only comparatively few, in particular Nile perch (Lates niloticus), various cichlids (especially tilapias) and clupeids (Limnothrissa miodon), have been successful in establishing themselves. Those that have become established have had obvious but unquantifiable impacts on the indigenous faunas.It is difficult to differentiate between the effects of fishing and of the presence of alien fish on the fish species composition of the lakes (Witte et al., 1992). Many of the lakes were overfished before introductions were made, with a resultant decline in some species, especially the larger ones, and the virtual disappearance of others. Some lake fish faunas, such as those of Lakes Kyoga and Victoria, which have been subjected to the perturbations described above, continue to change rapidly (Ogutu-Owayo, 1990b).There is a fundamental need to collect biological information on the fish communities of African lakes for effective management, resulting not only in the conservation of unique fish faunas but also the production of sustainable fish yields for the people relying on this source of protein. This information is required before any more introductions of exotic fish are made.     

Fish species and trophic diversity of haplochromine cichlids in the Kyoga satellite lakes (Uganda)

Prior to the 1980s, lakes Kyoga and Victoria previously supported an exceptionally diverse haplochromine fish fauna comprising at least 11 trophic groups. The species and trophic diversity in these lakes decreased when the introduced Nile perch depleted haplochromine stocks. From December 1996 to October 1998, we studied species and trophic diversity of haplochromine fishes in six satellite lakes without Nile perch in the Kyoga basin and compared them with the Kyoga main lake against historical data from Lake Victoria where Nile perch were introduced. Forty‐one species were found in the study area, of which, the Kyoga satellite lakes contributed 37 species in comparison to only 14 from the Kyoga main lake. Analysis of trophic diversity based on 24 species that contained food material revealed seven haplochromine trophic groups (insectivores, peadophages, piscivores, algal eaters, higher plant eaters, molluscivores and detritivores) in the Kyoga satellite lakes in comparison to two trophic groups (insectivores and molluscivores) in the Kyoga main lake. Many of the species and trophic groups of haplochromines depleted by the introduced Nile perch in lakes Kyoga and Victoria still survive in the Kyoga satellite lakes. This is attributed to the absence of Nile perch in those lakes. Nile perch has been prevented from spreading into the satellite lakes by swamp vegetation that separate them from the main lakes. If these swamps prevent Nile perch from spreading into the lakes, it is possible to conserve fish species, especially haplochromines, which are threatened by introduction of Nile perch in the main lakes.     

The taxonomy of East African Nile Perch, Lates spp. (Perciformes, Centropomidae)

The type specimens of Lates albertianus, L. macrophthalmus, L. niloticus rudolfianus and L. niloticus longispinis are contrasted with each other, with riverine L. niloticus niloticus and with a collection of Nile Perch from Lake Victoria. Contrary to current belief L. niloticus niloticus was found to differ both from L. albertianus and from L. niloticus rudolfianus , but no significant difference was found between L. albertianus and L. niloticus rudolfianus . The Lake Victoria collection was found to differ from all other taxa, suggesting that the characters currently used in the taxonomy of Lates are inappropriate for this purpose. It is recommended that a reappraisal of Nile Perch taxonomy be made using more modern techniques and that studies are initiated to discover how characters change during development under differing environmental conditions.     

Little evidence for morphological change in a resilient endemic species following the introduction of a novel predator

Human activities, such as species introductions, are dramatically and rapidly altering natural ecological processes and often result in novel selection regimes. To date, we still have a limited understanding of the extent to which such anthropogenic selection may be driving contemporary phenotypic change in natural populations. Here, we test whether the introduction of the piscivorous Nile perch, Lates niloticus, into East Africa's Lake Victoria and nearby lakes coincided with morphological change in one resilient native prey species, the cyprinid fish Rastrineobola argentea. Drawing on prior ecomorphological research, we predicted that this novel predator would select for increased allocation to the caudal region in R. argentea to enhance burst‐swimming performance and hence escape ability. To test this prediction, we compared body morphology of R. argentea across space (nine Ugandan lakes differing in Nile perch invasion history) and through time (before and after establishment of Nile perch in Lake Victoria). Spatial comparisons of contemporary populations only partially supported our predictions, with R. argentea from some invaded lakes having larger caudal regions and smaller heads compared to R. argentea from uninvaded lakes. There was no clear evidence of predator‐associated change in body shape over time in Lake Victoria. We conclude that R. argentea have not responded to the presence of Nile perch with consistent morphological changes and that other factors are driving observed patterns of body shape variation in R. argentea.     

Interactions Between Nile Perch, Lates niloticus, and Other Fishes in Lake Nabugabo, Uganda

The introduction of the predatory Nile perch, Lates niloticus, into the Lake Victoria basin coincided with a dramatic decline in fish species richness and diversity. This study focused on interactions between Nile perch and indigenous fishes in Lake Nabugabo, Uganda, a small satellite lake of Lake Victoria. We evaluated how the foraging impact of juvenile Nile perch on prey fishes varied with the size of the predator. We also evaluated the role of wetland ecotones in minimizing interaction between Nile perch and indigenous fishes. Wetland ecotones in Lake Nabugabo were characterized by complex structure (e.g., dense vegetation) and lower dissolved oxygen levels than non-wetland (exposed) areas. Nile perch (8.6–42.2cm, TL) were 3.7 times more abundant in offshore exposed areas than in inshore areas near wetland ecotones, and the proportion of Nile perch using wetland and exposed areas was independent of their body size. However, species richness was higher in waters at wetland ecotones than in exposed areas. Nile perch (5–35cm, TL) exhibited a shift in diet at approximately 30cm TL from feeding primarily on invertebrates to piscivory. Although the shift to piscivory occurred at approximately the same body size for Nile perch from both wetland and exposed habitats, the shift to piscivory was less abrupt in Nile perch captured near wetland ecotones. Nile perch from wetland areas consumed a greater diversity and a larger percentage of fish prey than those from exposed sites. However, the low abundance of Nile perch in wetland ecotones suggested that interaction between predator and prey in these areas is much reduced.