Genetic affinities of an introduced predator: Nile perch in Lake Victoria, East Africa

Although the introduction of Nile perch, Lates niloticus , to Lake Victoria has received intense global attention, especially in relation to its impact on endemic cichlid species and on fishery yields, fundamental information on its taxonomy and population genetics is lacking. Most importantly, the introduced fish originated from two lakes (Lakes Albert and Turkana) containing three Lates species, and it has never been entirely clear which of these became established in Lake Victoria, or indeed whether the Lake Victoria population is derived from hybridization between Lates species. In addition, genetic drift caused by the relatively small founder population (≈ 400), the initially slow population increase followed by a period of explosive population growth, and selection pressures in the new environment may have resulted in substantial genetic changes. Allozyme data indicated that the introduced Nile perch of Lake Victoria were mainly L. niloticus from Lake Albert, although maximum likelihood estimates of stock contributions (GSI) suggested the presence of L. macrophthalmus. In contrast, introduced Nile perch in adjacent smaller lakes (Lakes Kyoga and Nabugabo) appeared to be entirely L. niloticus . The effect of the introductions on allozyme diversity varied among lakes and appeared to be uncorrelated to the number of fish introduced.     

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 .     

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.     

The catfish fauna of Lake Victoria after the Nile perch upsurge

Stocks of the indigenous catfish species of Lake Victoria have decreased dramatically since the beginning of the 1980s. This decline coincided with the Nile perch boom and concomitant ecological changes in the lake. In deep water, where Nile perch densities were higher, the decline proceeded more quickly than in shallow water. In the former all catfishes eventually vanished. Of the two largest species, Clarias gariepinus and Bagrus docmak, juveniles disappeared faster than adults. This indicates that predation by Nile perch may have played an important role in their decline. Other possible impacts were the deoxygenation of deepwater areas and the decline of haplochromine cichlids which were an important food source for B. docmak, C. gariepinus and Schilbe intermedius. The various catfish species were not all affected to the same extent. The endemic Xenoclarias eupogon, which lived predominantly in deep water, may have become extinct. B. docmak currently seems to be mainly restricted to refugia in rocky habitats. Synodontis victoriae and S. afrofischeri are still present in small numbers in shallow littoral areas. Schilbe intermedius and C. gariepinus seem to be the least affected of the catfishes in littoral and sublittoral areas. This may be caused, among other reasons, by their smaller habitat overlap with Nile perch than the other species. S. intermedius is partly pelagic, and a considerable part of the C. gariepinus stock lives in bodies of water surrounding the lake. The patterns of decline of the catfishes are very similar to those observed for haplochromine cichlids in the lake. The importance of catfishes for the fisheries in the lake is currently negligible.     

Impact of the Nile perch on the fisheries of Lakes Victoria and Kyoga

The combined effects of lack of effective management, over-exploitation with destructive fishing gear and interspecific competition, particularly among tilapiines have had profound effects on the fish stocks of lakes Victoria and Kyoga. It has been proposed that these have been more important in the decline of the indigenous fisheries than predation or competition from Nile perch.     

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.     

Changes in relative abundance of zooplankton in northern Lake Victoria,East Africa

The zooplankton community of Lake Victoria was studied between March 1990 and February 1991 with the aim of identifying its constituent groups/species, their distribution, abundance, and long term changes in community structure. Zooplankton samples were taken from four stations using plankton nets (75–300 m mesh size) hauled vertically through the water column. The plankton comprised mainly Crustacea and to a lesser extent early stages of aquatic insects. Cyclopoid copepods, their nauplii and copepodites constituted the most frequent group at all stations. Calanoid copepods, Cladocera and Caridina nilotica (Roux), though widely distributed, contributed a small proportion. Chaoborus larvae and pupae and mites were occasionally present. A comparison of the relative proportions of the main zooplanktonic groups from three sources revealed remarkable changes in community structure since 1931. These changes are discussed with respect to predation, eutrophication and other changes in the food web structure of the lake.     

Ontogenetic shifts in phenotype–environment associations in Nile perch,Lates niloticus (Perciformes: Latidae) from Lake Nabugabo,Uganda

Habitat‐associated trait divergence may vary across ontogeny if there are strong size‐related shifts in selection pressures. We quantified patterns of phenotypic divergence in Nile perch (Lates niloticus) from ecologically distinct wetland edge and forest edge habitats in Lake Nabugabo, Uganda, and we compared patterns of divergence across three size classes to determine whether trends are consistent through Nile perch ontogeny. We predicted that inter‐habitat variation in biotic (e.g. vegetation structure) and abiotic (e.g. dissolved oxygen concentration) variables may create divergent selective regimes. We compared body morphology using geometric morphometrics and found substantial differences between habitats, although not all trends were consistent across size classes. The most striking aspects of divergence in small Nile perch were in mouth orientation, head size, and development of the caudal region. Medium‐sized Nile perch also showed differences in mouth orientation. Differences in large individuals were related to eye size and orientation, as well as caudal length. The observed patterns of divergence are consistent with functional morphological predictions for fish across divergent trophic regimes, high and low predation environments, and complex and simple habitats. Although this suggests adaptive divergence, the source of phenotypic variation is unknown and may reflect phenotypic plasticity and/or genetic differences. © 2013 The Linnean Society of London, Biological Journal of the Linnean Society, 2013, 110 , 449–465.     

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.     

Reproductive potential of the Nile perch,Lates niloticus L. and the establishment of the species in Lakes Kyoga and Victoria (East Africa)

The size at first maturity, sex ratio and fecundity of L. niloticus in Lake Kyoga have been examined, and compared with the situation in other aquatic systems. The species has the ability to reproduce enormously. It produces up to 16 million eggs. In Lake Kyoga, fecundity (F) increases with length (L) in cm according to the equation: % MathType!MTEF!2!1!+-% feaafiart1ev1aaatCvAUfeBSjuyZL2yd9gzLbvyNv2CaerbuLwBLn% hiov2DGi1BTfMBaeXatLxBI9gBaerbd9wDYLwzYbItLDharqqtubsr% 4rNCHbGeaGqiVu0Je9sqqrpepC0xbbL8F4rqqrFfpeea0xe9Lq-Jc9% vqaqpepm0xbba9pwe9Q8fs0-yqaqpepae9pg0FirpepeKkFr0xfr-x% fr-xb9adbaqaaeGaciGaaiaabeqaamaabaabaaGcbaGaciOraiaac2% dacaGGGaGaaiinaiaac6cacaGG0aGaai4maiaacAdacaGGGaGaaiiE% aiaacccacaGGXaGaaiimamaaCaaaleqabaGaaiylaiaacAdaaaGcca% GGmbWaaWbaaSqabeaacaGGYaGaaiOlaiaacMdacaGGYaaaaaaa!44D8!\[\operatorname{F} = 4.436 x 10^{ - 6} L^{2.92} \]Information from different habitats shows that females grow to a larger size than males but the growth rate is the same in both sexes. Males mature earlier than females at 50–65 cm total length with females maturing between 60–95 cm. There are about twice as many males as females. The rapid establishment of L. niloticus in Lake Kyoga and the Nyanza Gulf of Lake Victoria following its introduction is attributed to the high reproductive potential of the species under favourable environmental conditions.     

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     

Intrasexual competition among females and the stabilization of a conspicuous colour polymorphism in a Lake Victoria cichlid fish

The maintenance of colour polymorphisms within populations has been a long-standing interest in evolutionary ecology. African cichlid fish contain some of the most striking known cases of this phenomenon. Intrasexual selection can be negative frequency dependent when males bias aggression towards phenotypically similar rivals, stabilizing male colour polymorphisms. We propose that where females are territorial and competitive, aggression biases in females may also promote coexistence of female morphs. We studied a polymorphic population of the cichlid fish Neochromis omnicaeruleus from Lake Victoria, in which three distinct female colour morphs coexist: one plain brown and two blotched morphs. Using simulated intruder choice tests in the laboratory, we show that wild-caught females of each morph bias aggression towards females of their own morph, suggesting that females of all three morphs may have an advantage when their morph is locally the least abundant. This mechanism may contribute to the establishment and stabilization of colour polymorphisms. Next, by crossing the morphs, we generated sisters belonging to different colour morphs. We find no sign of aggression bias in these sisters, making pleiotropy unlikely to explain the association between colour and aggression bias in wild fish, which is maintained in the face of gene flow. We conclude that female-female aggression may be one important force for stabilizing colour polymorphism in cichlid fish.     

Phytoplankton community structure and environment in the Kenyan waters of Lake Victoria

• 1 Phytoplankton species composition, numerical abundance, spatial distribution and total biomass measured as chlorophyll a concentration were studied in relation to environmental factors in September 1994 (dry season) and March 1995 (rainy season), respectively, in the Kenyan waters of Lake Victoria; 103 species were recorded.
• 2 Blue‐green algae (Cyanophyceae) were most diverse, followed by diatoms (Bacillariophyceae), green algae (Chlorophyceae) and dinoflagellates (Dinophyceae).
• 3 Twinspan separated the phytoplankton communities in the Nyanza Gulf and those in the open lake during both seasons. During the dry season, the Nyanza Gulf was strongly dominated by blue‐greens, while diatoms dominated in the open lake. During the rainy season, blue‐greens remained dominant in the Nyanza Gulf although the number of species found was lower than during the dry season; in the open lake, blue‐greens replaced diatoms as the dominant group and there were more species than in the dry season.
• 4 Canonical correspondence analysis indicated that the phytoplankton species distribution was significantly correlated with turbidity during the dry season and with SiO₂ during the rainy season. Chlorophyll a concentrations ranging from 2.0 to 71.5 mg m^‐3 in the dry season and 2.0–17.2 mg m^‐3 in the rainy season confirm earlier reports of increasing phytoplankton biomass in Lake Victoria since the 1960s.

     

Potential fishery of Nile perch Lates niloticus Linne (Pisces: Centropomidae) in Nyanza Gulf of Lake Victoria,East Africa

From bottom trawls, Lates niloticus (Lin.) in Nyanza Gulf was observed to have a gulf-wide distribution. Lates dominated the catch at Homa point. Catch rates in kg h^-1 varied from one location to another and from month to month.Its estimated production is 11 000 metric tons, netting about 22 000 000 Kenya shillings (US $2 000 000).Abundance of food items such as freshwater shrimps of the genus Caridina, and Engraulicypris argenteus, have been used as basis for estimating the growth and production of Lates.     

Decline of the African lungfish (Protopterus aethiopicus) in Lake Victoria (East Africa)

Catch and effort data for the period 1973–1990 demonstrate a dramatic decline of lungfish in the Tanzanian waters of Lake Victoria. Bottom trawl catches in the Mwanza Gulf showed a decline in catch rates from 67.5 kg h⁻¹ in 1973 to 5.5 kg h⁻¹ in 1986. Trawling of commercial vessels in the Speke Gulf revealed a decline in lungfish catches from 1.3 kg h⁻¹ in 1986 to 0.07 kg h⁻¹ in 1990. The development of anoxia in the deeper waters of Lake Victoria, the algal blooms, and the decline of water transparency, all associated with eutrophication, are not likely to have contributed to the decreased catch rate. However, the lungfish decline may reflect the interaction of overexploitation by the fishery and a low level of Nile perch predation that restricts lungfish to wetland refugia. We suggest that this may have been reinforced over the past few decades by large-scale conversion of wetlands to agricultural land and harvesting of nest-guarding male lungfish leading to decreased recruitment of young.     

The Shrimp Caridina nilotica in Lake Victoria (East Africa), Before and After the Nile Perch Increase

The shrimp Caridina nilotica is a major prey of the introduced Nile perch in Lake Victoria. In spite of heavy predation, the density of shrimps increased after the Nile perch boom and the concomitant disappearance of the haplochromine cichlids. In the same period, the mean size of gravid shrimps and the size at first maturity declined. This seems to indicate an increased predation pressure on adult shrimps. Before the Nile perch upsurge, specialised shrimp eaters and piscivores, among the haplochromine cichlids, only took adult shrimps, whereas we assume that most haplochromines used to include juvenile shrimps into their diet. Another important predator on adult shrimps was Bagrus docmak. The combined density of predators on adult shrimps in the pre-Nile perch era was estimated at 10 kg ha⁻¹ and the potential predators on juveniles were estimated at 170 kg ha⁻¹. After the Nile perch upsurge, only Nile perch up to 10 cm TL and Rastrineobola argentea fed on juvenile shrimps (ca. 36 kg ha⁻¹) and Nile perch from 10 to 50 cm TL (ca. 13 kg ha⁻¹) fed on adults. These rough estimates of the biomass of predators on shrimps before and after the Nile perch upsurge indicate a reduced predation pressure on juvenile shrimps. The disappearance of the haplochromines may have released competition with small Nile perch for juvenile shrimps, thus enhancing the recruitment of Nile perch.     

PERMANENT GENETIC RESOURCES: Isolation and characterization of microsatellite loci in the cichlid fish in Lake Victoria, Haplochromis chilotes

Twelve short tandem repeat markers were successfully isolated from a cichlid, Haplochromis chilotes, in Lake Victoria, and characterized in Haplochromis pyrrhocephalus. The microsatellite regions of these markers were found to have between two and 48 alleles with heterozygosity ranging from 0.07 to 0.97. No loci showed significant departures from the Hardy–Weinberg or linkage equilibrium after the Bonferroni correction (P > 0.05). Cross‐species amplification in other cichlids of Lake Victoria, Haplochromis laparogramma, Lithochromis rubripinnis, L. rufus and Haplochromis sp. ‘rockkribensis’, was successful.     

Low genetic differentiation among morphologically distinct Labeobarbus species (Teleostei: Cyprinidae) in the Lake Victoria and Albertine basins,Uganda: insights from mitochondrial DNA

Labeobarbus altianalis and L. bynni bynni are hexaploid cyprinid fishes in the genus Labeobarbus. In the Great Lakes region of Africa, these two large-bodied barbs exhibit considerable morphological variations. Their intraspecific classification, currently based on geographical distribution and morphological variation, is of limited application due to the overlapping nature of the water systems. Patterns of genetic variation in the Albertine and Victoria drainage basins in Uganda were assessed using mitochondrial sequences of the control region (470 basepairs) and the cytochrome b gene (627 basepairs). At the control region 36 distinct haplotypes were observed in a total of 177 individuals. Relatively low overall nucleotide diversity (π = 0.010) was observed. Hierarchical analysis of the molecular variance revealed significant differentiation between populations in the total sample (F _ST = 0.172, p < 0.001), among populations within each basin (F _SC = 0.047, p < 0.001), but no significant subdivision among basins (F _CT = 0.131, p > 0.05). Phylogenetic analysis of the control region haplotypes resulted in a star-shaped phylogeny, with no clusters that reflect current taxonomic designation, morphotypes or basins. Phylogenetic analysis of cytochrome b gene sequences resulted in a tree topology similar to that observed for the control region. Thus the morphological characteristics separating the two species are intraspecific variation and the whole sample probably belongs to the same species.