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 and have been superseded by O. niloricus .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.     

The destruction of an endemic species flock: quantitative data on the decline of the haplochromine cichlids of Lake Victoria

Synopsis The Lake Victoria fish fauna included an endemic cichlid flock of more than 300 species. To boost fisheries, Nile perch (Lates sp.) was introduced into the lake in the 1950s. In the early 1980s an explosive increase of this predator was observed. Simultaneously, catches of haplochromines decreased. This paper describes the species composition of haplochromines in a research area in the Mwanza Gulf of Lake Victoria prior to the Nile perch upsurge. The decline of the haplochromines as a group and the decline of the number of species in various habitats in the Mwanza Gulf was monitored between 1979 and 1990. Of the 123+ species originally caught at a series of sampling stations ca. 80 had disappeared from the catches after 1986. In deepwater regions and in sub-littoral regions haplochromine catches decreased to virtually zero after the Nile perch boom. Haplochromines were still caught in the littoral regions where Nile perch densities were lower. However, a considerable decrease of species occurred in these regions too. It is expected that a remnant of the original haplochromine fauna will survive in the littoral region of the lake. Extrapolation of the data of the Mwanza Gulf to the entire lake would imply that approximately 200 of the 300+ endemic haplochromine species have already disappeared, or are threatened with extinction. Although fishing had an impact on the haplochromine stocks, the main cause of their decline was predation by Nile perch. The speed of decline differed between species and appeared to depend on their abundance and size, and on the degree of habitat overlap with Nile perch. Since the Nile perch upsurge, the food web of Lake Victoria has changed considerably and the total yield of the fishery has increased three to four times. Dramatic declines of native species have also been observed in other lakes as a result of the introduction of alien predators. However, such data concern less speciose communities and, in most cases, the actual process of extinction has not been monitored.     

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.     

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 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.     

Nile perch and the transformation of Lake Victoria

The transformation of Lake Victoria that began in 1980 followed the population explosion of Nile perch Lates niloticus, causing the apparent extirpation of 500+ endemic haplochromine species and dramatic physico-chemical changes. Officially introduced in 1962–1963, but present earlier, the reasons for the long delay before its population exploded are discussed. The hypothesis that it occurred only after the haplochromine decline is evaluated, but haplochromines declined only after the Nile perch expansion began. The sudden eutrophication of the lake was attributed to Nile perch, but evidence of eutrophication from 1950 onwards led some researchers to conclude that it was the result of climatic changes. We conclude that the haplochromine destruction disrupted the complex food webs that existed prior to the upsurge of Nile perch. The depletion of fish biomass by Nile perch may have been the source of extra phosphorus responsible for the eutrophication of the lake. After the Nile perch explosion in 1980 the fish population came to be dominated by only three species, but fisheries productivity increased at least 10-fold. Fishing has caused demographic changes in Nile perch, which may have allowed some haplochromine species to recover. The condition of the lake appears to have stabilised since 2000, partly because the fish biomass has risen to at least 2 × 10⁶ t, replacing the ‘lost’ biomass and restoring some ecosystem functioning.     

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.     

The invasion of an introduced predator,Nile perch (<Emphasis Type="Italic">Lates niloticus</Emphasis>, L.) in Lake Victoria (East Africa): chronology and causes

Nile perch, a large predatory fish, was introduced into Lake Victoria in 1954. The upsurge of Nile perch in Lake Victoria was first observed in the Nyanza Gulf, Kenya, in 1979. In Ugandan waters this occurred 2–3 years later and in the Tanzanian Mwanza Gulf 4–5 years later. At the beginning of the upsurge in the Mwanza Gulf in 1983/1984 only sub-adult and adult fishes were found. The first juveniles appeared in 1985, suggesting that the initial increase of Nile perch was mainly caused by migration of sub-adults and adults. Shortly after the onset of trawl fishery in the area in 1973, haplochromines in the Mwanza Gulf started to decline. The final disappearance of the haplochromines, in 1987, only occurred after the Nile perch boom, and despite the abandoning of the haplochromine fishery in 1986. We hypothesize that the decline of haplochromines decreased predation on and competition with juvenile Nile perch and then facilitated survival of these juveniles. Consequently the immigration of sub-adult and adult Nile perch in an area may have paved the way for successful recruitment. Over-exploitation of haplochromine cichlids in the 1970s in the Nyanza Gulf, where the Nile perch upsurge was first observed, may have played a similar role.     

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 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.     

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.     

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.     

Biological diversity of the Yala Swamp lakes, with special emphasis on fish species composition, in relation to changes in the Lake Victoria Basin (Kenya): threats and conservation measures

During the second half of the last century, the Lake Victoria ecosystem has undergone drastic ecological changes. Most notable has been the decline in the populations of many endemic cichlid fishes. The lake has lost nearly 200 haplochromines and one tilapiine, Oreochromis esculentus. The above changes have been attributed to effects of species stocking and, in particular, from predation pressure by the introduced Nile perch, Lates niloticus. Other factors that have led to the decline of the endemic species include intensive non-selective fishing, extreme changes in the drainage basin, increased eutrophication, and the invasion of the lake by the water hyacinth, Eichhornia crassipes. However, the remnants of some species that had disappeared from Lake Victoria occur abundantly in the Yala Swamp lakes (Kanyaboli, Sare and Namboyo). This paper discusses the biodiversity of the swamp and the three lakes and gives suggestions for their conservation.     

Changes in the diet of Nile perch,Lates niloticus (L), in the Mwanza Gulf,Lake Victoria

Samples of Nile perch (Lates niloticus L.) were collected for stomach analysis from trawl catches conducted in the Mwanza Gulf from September 1986 to September 1988. Initially haplochromine cichlids formed the main food item for the Nile perch. Despite their decline, haplochromines still formed the major part of its diet in 1986. After the virtually complete disappearance of the haplochromines in 1987 and 1988, the benthic shrimp Caridina nilotica, the pelagic cyprinid, Rastrineobola argentea and juvenile Nile perch became the main food of Nile perch. Twenty-four trawl sessions were conducted to determine the relative quantities of respective prey taken. When present, Caridina is the major prey, mainly taken by day. If absent, juvenile Nile perch constitute the main part of the diet. Seasonal differences in diet composition probably reflect seasonal fluctuations in the abundance of the main prey species.     

Effects of Nile perch (Lates niloticus) introduction into Lake Victoria,East Africa,on the diet of Pied Kingfishers (Ceryle rudis)

In recent years the ichthyofauna of Lake Victoria, the world's largest tropical lake, has gone through dramatic changes. The population of Nile perch, a large predator which has been introduced into the lake by man, increased explosively at the expense of many haplochromine cichlid species. At the same time, numbers of a small cyprinid (dagaa) rose sharply.

Previously Pied Kingfishers on Lake Victoria fed mainly on haplochromines. Only the youngest nestlings depended on dagaa as primary food. The current diet of adult birds clearly reflects the changes which have occurred in the fish community. Pellet analysis reveals a shift towards a diet composed of almost 100% dagaa.

The change in prey species composition has increased the number of fish a kingfisher needs to catch daily in order to meet its energetic demands, because:

1. (1)

   the mean size of haplochromines is larger than that of dagaa;

2. (2)

   (2) the mean size of dagaa has decreased since the increase in Nile perch;

3. (3)

   (3) the weight of dagaa is lower than that of haplochromines of equal size;

4. (4)

   (4) mainly juvenile dagaa and adults in poor condition are accessible to kingfishers.

     

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 .     

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.     

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.