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.     

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.     

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.     

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.     

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.     

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.     

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.     

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.     

Spatial and seasonal patterns in the feeding habits of juvenile Lates niloticus (L.), in the Mwanza Gulf of Lake Victoria

Flexibility in the feeding habits of juvenile Nile perch (1–30 cm total length) was studied from September 1988 to September 1989 at four sites (depth range: 1–25 m) in the Mwanza Gulf of Lake Victoria. During this period haplochromine cichlids were virtually absent in the area. We looked at the combined effects of predator size, season and habitat. Stomach content analysis showed that with increase in size, the diet of Nile perch shifted from zooplankton and midge larvae, to macro-invertebrates (shrimps and dragonfly nymphs) and fish. At a size of 3–4 cm Nile perch shifted from size-selective predation on the largest cyclopoids to predation on the largest, less abundant, calanoids. Zooplanktivory ended at a size of ca. 5 cm. Although an ontogenetic shift in the diet of juvenile Nile perch was obvious at all sampling stations, the contribution of prey types appeared to be habitat related. With increasing water depth the frequency of occurrence in the diet of most prey types decreased, but that of shrimps increased. At the entrance of the gulf (20–25 m deep) shrimps were the main food source throughout the year. Halfway the gulf (12–16 m), Nile perch showed seasonality in their feeding behaviour. Shrimps were taken there especially during the rainy season (January to May) when their densities at this station were high, whereas cannibalism prevailed during the rest of the year. In an environment with Nile perch and dagaa as alternative prey, shrimps were taken almost exclusively. They could be regarded as a key prey for Nile perch between 5 and 30 cm.     

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     

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.

     

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 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.     

Diel vertical migration of major fish-species in Lake Victoria, East Africa

Understanding of migration patterns is essential in the interpretation of hydro-acoustic stock assessment data of partly demersal partly pelagic fish stocks. In this paper we provide this kind of information for some species that were common in the Mwanza Gulf of Lake Victoria in the 1980s, before and after the upsurge of introduced Nile perch (Lates niloticus). Detritivorous haplochromines and Nile tilapia (Oreochromis niloticus), both stay near the bottom during day and night. Feeding seems to occur predominantly during the day. The zooplanktivorous haplochromines and dagaa (Rastrineobola argentea) dwell near the bottom by day and migrate towards the surface during the night. They seem to follow their prey, zooplankton and lake-fly larvae. Piscivorous nembe (Schilbe intermedius) show similar migration patterns to zooplanktivorous fishes, but their behaviour cannot be unambiguously explained by pursuit of prey. Nile perch to some extend migrate into the column at night, though the majority remains near the bottom. Feeding takes place during day and night.     

Adaptive responses in resurgent Lake Victoria cichlids over the past 30 years

Textbook examples of adaptive radiation like the Galapagos finches and the East-African cichlids form a subject of major interest in evolutionary biology. Many of these species often show rapid morphological changes in response to a perturbed environment. The dramatic ecological changes in Lake Victoria during the past three decades, e.g. Nile perch predation and eutrophication, provide a unique opportunity to study environmental effects on cichlid morphology. Preliminary research has revealed that the lake’s haplochromines tend to be extremely plastic and sensitive to these environmental changes. So far, long-term ecomorphological studies at short-term intervals are extremely rare. In this study, we examined morphological changes over a 30 year period in six haplochromine species. Geometric morphometric analyses at intervals of approximately 3 years revealed adaptive responses. Three out of four resurgent haplochromines had a smaller head surface/caudal peduncle area (HS/CPA) ratio during the upsurge of the predatory Nile perch. During the same period, all four resurgent species had a larger cheek depth and a smaller eye size. The smaller HS/CPA ratio and larger cheek depth are likely to be adaptive responses to a high predation pressure and a diet shift to larger prey. The smaller eye size seems to be the result of a trade off between the eyes and other morphological structures in the smaller head of these species. Interestingly, the direction of the morphological changes was different between the four resurgent cichlid species and two species that became extremely rare or even may have gone extinct. The HS/CPA ratio increased in the extinct species where it decreased in the resurgent species. This study suggests that predation is a major driver of these morphological changes, which may be due to either phenotypic plasticity or adaptive changes.     

Food web structure and mercury transfer in two contrasting Ugandan highland crater lakes (East Africa)

Volcanic crater lakes scattered throughout western Uganda are important local sources of water and fish. Two representative but contrasting crater lakes near the Kibale National Park were sampled in 2000; the hyper‐eutrophic Lake Saka, which is highly affected by agricultural practices, and the mesotrophic Lake Nkuruba that is still surrounded by intact forest. The food web structures in these two lakes were assessed using stable nitrogen (δ¹⁵N) and carbon (δ¹³C) isotope analyses, and the mercury (THg) transfer patterns were quantified. The δ¹⁵N results indicate that food webs in both lakes are abbreviated, with only one to two trophic levels from primary consumers. The Lake Saka biota had distinctively enriched δ¹³C values compared with those in Lake Nkuruba, which may be due to ¹²C‐limited phytoplankton blooms in this lake. In Lake Nkuruba, two introduced tilapiine species and the introduced guppy Poecilia reticulata fed predominantly upon invertebrates and decomposed terrestrial plant material. In Lake Saka, the introduced Nile perch Lates niloticus appeared to occupy the top trophic position, but stable isotope values of the endemic haplochromine cichlids exclude those as Nile perch prey items. THg was found to biomagnify through the food web, reaching highest concentrations in P. reticulata in Nkuruba, which tended to be higher than for L. niloticus in Saka, suggesting increased bioavailability of THg in Nkuruba. Maximum THg concentrations in fish never approached WHO recommended guidelines (200 ng g^?1) designed to protect at‐risk groups.     

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.     

Feeding behaviour determining differential capture success of evasive prey in underyearling European perch (<Emphasis Type="Italic">Perca fluviatilis</Emphasis> L.) and roach (<Emphasis Type="Italic">Rutilus rutilus</Emphasis> (L.))

The effect of feeding behaviour on the prey capture efficiency of young-of-the-year European perch and roach was investigated in laboratory experiments using planktonic crustaceans possessing different escape abilities—Daphnia sp. and Cyclops sp. Two sets of experiments were performed. In the first set, the feeding efficiency and behaviour of 270 fish individuals were determined by stomach content analyses and video record evaluations. In the second set of experiments, analysis of attack-effort, which was evaluated as attack-distance and repeated strikes, was undertaken. Except for situations in which Daphnia was offered at high densities, the feeding efficiency of perch was significantly higher compared to roach in all other combinations of prey types and densities. Roach consumed significantly less prey compared to perch when feeding exclusively on the evasive Cyclops and when it was offered in a 1:1 ratio mixture with Daphnia. The mean swimming speed was similar in both fish species, but behavioural differences were evident during prey search and capture. Perch swam through the aquaria in short and fast movements that were interrupted by many stops. Roach exhibited rather continuous swimming that was punctuated by slowdowns instead of stops. The perch attacks were very intensive and repeated strikes occurred, particularly when feeding on evasive Cyclops. On the other hand, roach revealed strong schooling behaviour restricting the fish during inspection of the experimental aquaria. The distinct differences in feeding efficiency between perch and roach were demonstrated to be closely related to differences in their feeding behaviour. Discontinuous searching for prey, vigorous attacks, occurrence of repeated strikes and the absence of schooling increased perch prey capture efficiency, particularly when foraging on evasive copepods.     

Age and spread of the haplochromine cichlid fishes in Africa

The Haplochromini are by far the most species-rich cichlid fish tribe that originated along with the so-called primary radiation of the Lake Tanganyika cichlid species flock, i.e. at the same time during which the majority of the endemic Lake Tanganyika cichlid tribes emerged. Unlike the other tribes, the haplochromines are not restricted to Lake Tanganyika but distributed throughout Africa, except for the northwestern part of the continent. Haplochromine cichlids seeded the adaptive radiation of cichlid fishes in Lakes Malawi, Kivu, Victoria, Turkana, as well as in the now extinct paleo-Lake Makgadikgadi. Here we present a comprehensive phylogenetic and phylogeographic analysis of haplochromine cichlids that is based upon DNA sequences of two mitochondrial gene segments of riverine taxa covering all major African biogeographic regions where haplochromines are found. Our analysis revealed that six lineages of haplochromines originated within a short period of time, about 5.3-4.4 MYA. These haplochromine lineages show a highly complex phylogeographic pattern, probably severely influenced by climate- and/or geology-induced changes of the environment, with river capture events most likely playing an important role for species dispersal.