Nile perch (Lates niloticus,L.) and cichlids (Haplochromis spp.) in Lake Victoria: could prey mortality promote invasion of its predator?

The invasion of Nile perch into Lake Victoria is one of the iconic examples of the destructive effect of an introduced species on an ecosystem but no convincing explanation exists of why Nile perch only increased dramatically after a 25 year lag. Here, we consider this problem using a mathematical model that takes into account interactions between Nile perch and its cichlid prey. We examined competing hypotheses to explain Nile perch invasion and show that suppression of juvenile Nile perch by cichlids may cause the system to have two alternative stable states: one with only cichlids and one with coexistence of cichlids and Nile perch. Without cichlid predation on Nile perch, alternative stable states did not occur. Our analysis indicates that cichlid mortality, for example fishing mortality, may have induced the observed shift between the states.     

Predation by Nile perch Lates niloticus (L.) on Oreochromis niloticus (L.), Cyprinus carpio (L.), Mugil sp. and its role in controlling tilapia recruitment in Egypt

Vulnerability of three fish groups (tilapia, common carp, mullet) to predation by Nile perch Lates niloticus was evaluated. The study was conducted in aquaria and in cages placed in concrete ponds. Common carp Cyprinus curpio was the more vulnerable to predation by Nile perch followed in order by mullet ( Mugil cephulus and M. cupito ) and Nile tilapia Oreochromis niloricus . Medium-size Nile perch (30–32 cm t.l. ) were able to consume tilapia up to 14 cm t.l. , while tilapia of 15 cm total length were not eaten. Ratios of 1 : 10 to 1 : 15 between Nile perch and ready-to-spawn females of O. niloticus were found to be adequate to control tilapia reproduction.     

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.     

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 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.     

Dwelling at the oxycline: does increased stratification provide a predation refugium for the Lake Victoria sardine Rastrineobola argentea?

1. We tested the hypothesis that increased hypoxia in Lake Victoria provides a refugium for the cyprinid dagaa ( Rastrineobola argentea ) against hypoxia-sensitive Nile perch ( Lates niloticus ).
2. Hypoxia in the main habitat of dagaa was rare during 1979–80, but lasted 3–5 months in 1987 and 1988, during which time adult dagaa spent the day just above the oxycline instead of near the bottom.
3. Dissolved oxygen concentrations in the layer just above the oxycline were not critical for Nile perch, but a mass kill of this species following a sudden upwelling of hypoxic water suggested that oxycline-dwelling is risky.
4. Our data suggest no difference between dagaa and Nile perch in the level at which dissolved oxygen starts to limit their vertical distribution in the water column, but dagaa seems to be even more sensitive to extreme hypoxia (<1–2 mg O2 L−1) than Nile perch.
5. We argue that oxycline-dwelling dagaa are not seeking a predation refugium but that they are limited by low oxygen levels in reaching their feeding areas near the bottom.     

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     

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.     

Was Lates Late? A Null Model for the Nile Perch Boom in Lake Victoria

Nile perch (Lates niloticus) suddenly invaded Lake Victoria between 1979 and 1987, 25 years after its introduction in the Ugandan side of the lake. Nile perch then replaced the native fish diversity and irreversibly altered the ecosystem and its role to lakeshore societies: it is now a prised export product that supports millions of livelihoods. The delay in the Nile perch boom led to a hunt for triggers of the sudden boom and generated several hypotheses regarding its growth at low abundances – all hypotheses having important implications for the management of Nile perch stocks. We use logistic growth as a parsimonious null model to predict when the Nile perch invasion should have been expected, given its growth rate, initial stock size and introduction year. We find the first exponential growth phase can explain the timing of the perch boom at the scale of Lake Victoria, suggesting that complex mechanisms are not necessary to explain the Nile perch invasion or its timing. However, the boom started in Kenya before Uganda, indicating perhaps that Allee effects act at smaller scales than that of the whole Lake. The Nile perch invasion of other lakes indicates that habitat differences may also have an effect on invasion success. Our results suggest there is probably no single management strategy applicable to the whole lake that would lead to both efficient and sustainable exploitation of its resources.     

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.     

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.     

Alternative antipredator responses of two coexisting Daphnia species to negative size selection by YOY perch

Our study showed for the first time in nature that two coexistingDaphnia adopted alternative life history and behavioural strategiesto cope with negative size-selection predation by gape-limitedyoung-of-the-year (YOY) perch. We evaluated the phenotypic plasticityin life history and behavioural traits of two coexisting Daphniaspecies, D. pulicaria (2 mm) and D. galeata mendotae (1.4 mm),in response to seasonal changes in predation by YOY yellow perch(Perca flavescens) in a mesotrophic lake. We expected that thelarge-sized D. pulicaria, the most likely subjected to size-selectivepredation by YOY perch, will show stronger antipredator responsesthan the small-sized D. galeata mendotae. To test this hypothesis,we examined changes in life history and behavioural traits injuveniles and adults of both species during four YOY fish predationperiods that were selected based on the presence of YOY perchin the pelagic zone and the relative abundance of Daphnia preyin their gut contents. Our study supports the scenario of negativesize-selective predation by gape-limited YOY perch on both Daphniaspecies. The electivity index indicated that no daphnids witha body length > 1.75 mm were predated by YOY yellow perch.Coexisting Daphnia exhibited phenotypic plasticity in theirantipredator defenses based on their vulnerability to seasonalchanges in size-selective predation of YOY perch. Juvenile Daphniawere the targeted prey and they responded by a decreased bodylength. Behavioural defenses were the dominant strategy usedby both adult Daphnia populations to withstand high predation.A decreased size at maturity was not employed by Daphnia, exceptat the very end of the predation period. Behavioural defensesare short-term strategy adopted to avoid predation. Both antipredatordefenses became unnecessary expenses and were no longer sustainedafter the predation period.     

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 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.     

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.     

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.     

Intraspecific variation in gill morphology of juvenile Nile perch, Lates niloticus, in Lake Nabugabo, Uganda

Several studies have demonstrated intraspecific variation in fish gill size that relates to variation in dissolved oxygen (DO) availability across habitats. In Lake Nabugabo, East Africa, ecological change over the past 12 years has coincided with a shift in the distribution of introduced Nile perch such that a larger proportion of the population now inhabits waters in or near wetland ecotones where DO is lower than in open waters of the lake. In this study, we compared gill size of juvenile Nile perch between wetland and exposed (open-water) habitats of Lake Nabugabo in 2007, as well as between Nile perch collected in 1996 and 2007. For Nile perch of Lake Nabugabo [<20 cm total length (TL)], there was a significant habitat effect on some gill traits. In general, fish from wetland habitats were characterized by a longer total gill filament length and average gill filament length than conspecifics from exposed habitats. Nile perch collected from wetland areas in 2007 had significantly larger gills (total gill filament length) than Nile perch collected in 1996, but there was no difference detected between Nile perch collected from exposed sites in 2007 and conspecifics collected in 1996.     

The Lake Victoria environment: Its fisheries and wetlands — a review

Recent ecological changes in the Lake Victoria ecosystem have been attributed to the effects of species stockings and, in particular, from predation pressure by the Nile perch. Evidence for the decline of haplochromines due to predation by the Nile perch, while overwhelming, does not necessarily account for these gross ecological changes. Ecological theory predicts that natural fluctuation would occur in a fisheries where predator and prey species could alternate in abundance. The absence of a substantial recovery of the endemic species within cyclical abundance patterns in Lake Victoria (even though they could be delayed), particularly in the pelagic and profundal zones, points to other causal factors.Ecological changes have occurred in Lake Victoria since the turn of the century when modern fishing methods and techniques were introduced. As the human population increased and the catchment became more exposed to diverse socio-economic activities, further pressure on the ecological functioning of the lake was compounded by exotic species stockings. Thus, declining fisheries, wetland degradation and eutrophication are part of gross environmental changes that are likely to become more manifest with the increasing impact of the water hyacinth. It therefore seems appropriate to consider the ecological changes at both temporal and spatial scales and to re-examine some of the paradigms for ecological change.Although cataclysmic impacts may have occurred between the 1960's and the early 1980's, the basic cause of ecological changes, at least in the inshore zone, appears to result from human activities partly associated with a degradation of the riparian wetlands that exposes the aquatic ecosystem to catchment activities. Overexploitation and a reduction in habitat quality and quantity in this zone could be major factors in the ecological transformations. However, the importance of the littoral zone and the ecological impact of the Nile tilapia, another stocked species, are hardly known. An additional hypothesis is therefore put forward to determine whether or not wetland vegetation types in Lake Victoria are important fish habitats. This hypothesis allows for testing several interrelated sub-hypotheses about the relationships between wetland vegetation and the ecology of Nile tilapia. The water hyacinth, a recent feature of the shoreline and an additional strain to the already endangered buffer zone, may have to be regarded as a dynamic, but permanent, part of most of the littoral zone.Corresponding Editor: Prof. P. Denny