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

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

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

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

The food and feeding habits of Eutropius niloticus (Ruppell), family Schilbeidae (Osteichthyes: Siluriformes) in Lake Kainji,Nigeria

Studies on the food and feeding habits of Eutropius niloticus in Lake Kainji, Nigeria revealed that the fish is omnivorous but fed more on insects than on any other food. The most important insects were Povilla adusta Navas nymphs and imagos and adults of the common grasshoppers. Fish, particularly the clupeids Sierrathrissa leonensis Thys Van den Audernaerde and Pellonula afzeliusi Johnels, also formed an important part of the diet. There was seasonal variation of organisms selected for food and there were variations in the dietary items selected by different age groups. Accumulation of fat was found to correlate with seasonal feeding intensity. E. niloticus serves as food for other fishes.     

Fatty acid composition of muscle and heart tissue of Nile perch,Lates niloticus,and Nile tilapia,Oreochromis niloticus,from various populations in Lakes Victoria and Kioga,Uganda

The fatty acid composition in the heart tissue and muscle tissue of the Nile perch, Lates niloticus, and Nile tilapia, Oreochromis niloticus populations from Lakes Kioga and Victoria was determined by methanolysis and gas chromatography of the resulting fatty acid methyl esters. The analytical data were treated by multivariate principal component analysis. The most abundant individual fatty acids were palmitic acid (16:0), stearic acid (18:0), oleic acid (18:1n9), vaccenic acid (18:1n7), arachidonic acid (20:4n6) and docosahexaenoic acid (22:6n3). Due to high levels of both n6 and n3 fatty acids, the ratios of n3 to n6 were between 1 and 2, typical for freshwater fish species. Two Lake Victoria and one Lake Kioga populations of Nile tilapia and Nile perch were distinguished by the fatty acid profiles in their heart and muscle tissue. The heart tissue showed better separation than muscle tissue, due to dominance of polar phospholipids. It is rationalised that genetics are more important than diet in determining the fatty acid composition of the tissues.     

Comparative age and growth of Nile tilapia (<Emphasis Type="Italic">Oreochromis niloticus</Emphasis> L.) in lakes Nabugabo and Wamala,Uganda

Age and growth of Nile tilapia (Oreochromis niloticus) from Lake Nabugabo and Lake Wamala, Uganda, were determined using cross-sectioned sagittal otoliths. Marginal-increment and edge analyses of Nile tilapia otoliths from Lake Nabugabo indicated formation of two annuli per 12-month period. Opaque zones associated with faster growth were observed between April and June and between September and December, coincident with the two rainy seasons of the year. Within both lakes, males were larger at age than females. Nile tilapia from Lake Nabugabo, however, had faster growth rates than Nile tilapia from Lake Wamala, and fish >3 years old from Lake Nabugabo were larger at age than those from Lake Wamala. Ages ranged from 0 to 8.0 years for Nile tilapia from Lake Nabugabo, and from 0.5 to 6.5 years for tilapia from Lake Wamala. Differences in the patterns of growth in Nile tilapia between lakes may reflect, at least in part, the relatively energy-rich omnivorous diet of Nile tilapia in Lake Nabugabo versus a phytoplanktivorous diet in Lake Wamala. Diet differences of Nile tilapia between the two lakes are ascribed to trophic changes in the lakes due to the introduction of Nile perch (Lates niloticus) into Lake Nabugabo but not Lake Wamala. Alternatively, the greater exploitation of Nile tilapia in Lake Nabugabo may have resulted in increased growth rates, whereas Nile tilapia in Lake Wamala may be subject to slower, density-dependent growth. Handling editor: J. Cambray     

The evolution of the fishery of Oreochromis niloticus (Pisces: Cichlidae) in Lake Victoria

The Lake Victoria ecosystem has experienced changes associated with fishing levels, a rise in lake level in the 1960s, fish introductions, and human activities in the drainage basin. Following the fish introductions of the 1950s and early 1960s, Oreochromis niloticus has become the most abundant and commercially important species among the tilapiines, and the only species which has managed to co-exist with the Nile perch in Lakes Victoria and Kyoga. There is, however, little published information on the biology and ecology of the specie in the new habitats. It has therefore been found necessary to initiate studies on the characteristics of O. niloticus in Lake Victoria.     

Microcystin concentrations in Nile tilapia (<Emphasis Type="Italic">Oreochromis niloticus</Emphasis>) caught from Murchison Bay,Lake Victoria and Lake Mburo: Uganda

Catches of the economically important Nile tilapia (Oreochromis niloticus L.) from two eutrophic tropical lakes in Uganda, Lake Mburo and Murchison Bay, Lake Victoria, were examined to determine the presence of microcystins (MCs) in gut, liver and muscle of the fish. Analysis for MCs (RR, LR and YR) in both fish and water samples was by liquid chromatography coupled with mass spectroscopy (LC-MS) method. Physico-chemical parameters were also measured to establish the status of both lakes. MC-RR was the most prominent MC detected in Lake Mburo and Murchison Bay samples, there was no evidence of significant seasonal variation in the concentration of MCs in fish tissue. MCs were detected in all water samples from both study lakes. The mean concentration of MCs in water was found higher in dry times for Lake Mburo (P < 0.05) and higher in wet times for Murchison Bay (P < 0.05). MC concentrations in the fish guts were positively related with MC concentrations in water samples from Murchison bay (P < 0.05), no such correlation was found in Lake Mburo. In eutrophic tropical lakes, fish seem to have a high tolerance to the toxicity of cyanotoxins including MCs. However, there is a possibility of accumulating these toxins in their tissue with the threat of transferring them higher up in the food chain. Due to a low sample size and short sampling period, the results can only serve to highlight the potential risk of MC accumulation in Nile tilapia in these lakes. Further studies are needed for the purpose of risk assessment.     

Some biological aspects and life history strategies of Nile tilapia Oreochromis niloticus (L.) in Lake Victoria, Kenya

The life history characteristics of introduced Nile tilapia (Oreochromis niloticus L.) in Lake Victoria, including, sex ratio, fecundity, reproduction, weight‐length relationship and body condition were studied and compared with those of other populations. Samples were collected by trawling and seining in the Kenyan sector of Lake Victoria between June 1998 and December 2000. Males predominated over females (sex ratio 1.42 : 1 : 00). O. niloticus spawned throughout the year but with a peak between December and June. Length at first maturity was (mean ± SD) 30.81 ± 0.09 for females and 34.5 ± 6 0.48 for males. There was little seasonal variation in relative condition, which ranged from 0.92 to 1.05 in males and 0.94 to 1.07 in females. Gonadosomatic index (GSI) was low during the postspawning period (July to October) and high during the protracted breeding period (December and June). Fecundity ranged from 905 to 7619 oocytes for fish of 28 to 51 cm total length (TL) respectively. The relationships between fecundity (F) and total length (L), weight (W) and ovary weight (OW) were: F = 8.159L^1.53, F = 96.269W^0.4504, F = 1806 + 39.4OW. The slope b of the weight‐length relationship was 3.08–3.32 for males and 3.07–3.22 for females. Growth was allometric in both cases and was significantly different from the expected value of 3. The life history strategy of O. niloticus is discussed in context of environmental changes occurring in the lake.     

The food habits of an herbivorous fish (Oreochromis niloticus Linn.) in Lake Awasa,Ethiopia

The natural food ofOreochromis niloticus in Lake Awasa was studied from the stomach contents of 10 fish measuring 18–32 cm (standard length) monthly from August 1984 to April 1986.Chroococcus, Oscillatoria andBotryococcus were found to be important food items in the diet. Animal foods were observed on rare occasions and these were mainly Rotifera. Blue-green algae as a group contribute about 28.1% of the ash free dry weight (AFDW) whereas diatoms and green algae (excludingBotryococcus) make 2.31% and 2.39% of AFDW, respectively.Botryococcus, detritus and other algae comprised 67.2% of AFDW and it was suspected that more than half of this was contributed byBotryococcus. Blue-green algae, which were especially abundant in the diet during the dry season, November–February, are nutritionally most important. Seasonal variation in algal species composition could influence the quality of food ingested by the fish.     

Trophic interrelationships between the exotic Nile tilapia, <Emphasis Type="Italic">Oreochromis niloticus</Emphasis> and indigenous tilapiine cichlids in a subtropical African river system (Limpopo River,South Africa)

The stable isotope ratio and seasonal changes in diet of two indigenous (Oreochromis mossambicus, Tilapia rendalli) and one exotic (Oreochromis niloticus) tilapiine cichlids in the subtropical Limpopo River, South Africa were investigated to determine patterns of resource partitioning. Stomach contents of O. niloticus and O. mossambicus indicated high dietary overlap across size class, habitat and season, with both species primarily feeding on vegetative detritus. However, stable isotope analysis revealed that the two Oreochromis species had different stable isotope ratios derived from different food sources. The relatively δ¹³C-depleted O. niloticus indicates a phytoplankton-based diet, while the δ¹³C-enriched O. mossambicus indicates a macrophagous diet dominated by vegetative detritus and periphyton. The high similarity in stomach contents and the interspecific differences in isotopic composition reveal fine-scale patterns of food resource partitioning that could be achieved through selective feeding. Tilapia rendalli was largely macrophagous and fed mainly on aquatic macrophytes and had a low dietary overlap with both O. niloticus and O. mossambicus. In the Limpopo River, detritus and algae are probably the most abundant food resources and the causal factors responsible for the observed patterns of resource partitioning among the tilapiines are usually difficult to ascertain. Fish may be able to perceive food resources in terms of the dynamics that determine their availability. Detailed studies of variation in food resource availability and fish habitat use within the system are needed to evaluate this hypothesis.     

Feeding ecology of Oreochromis niloticus (Linnaeus) & Tilapia Zillii (Gervias) in a Nile canal

Fish stomachs, 1149 for Tilapia zillii and 1698 for Oreochromis niloticus, were collected from April 1984 till April 1985 from a Nile canal in the Egyptian delta. Both species showed a significant overlap of diet (Schoener's index) among individuals ranging from 12 to 17.9 cm standard length. Outside this range differences in the importance of food of animal origin occurred. Macrophytes were the main food of O. niloticus and aquatic insects of the food of T. zillii. This initial diet overlap may have contributed to irregularity in the increase of full stomachs with increasing length, and to a decline of the stomach index (SI = stomach weight × 100/fish weight) with increase in length in O. niloticus. The stomach of either species correlated significantly with water temperature, but with daily photoperiod in O. niloticus only.     

Studies on the biology of Nile tilapia (Oreochromis niloticus) in Lake Victoria,Kenya: in light of intense fishing pressure

Nile tilapia Oreochromis niloticus is now the most abundant and commercially important tilapiine in Lake Victoria. From the total of 1 512 fish sampled from commercial gill net fisheries during 2014 and 2015, 809 (54%) were males and 672 (44%) were females, giving an overall sex ratio of 1.20 males: 1.00 females. The mean (± SE) length and weight for all fish were 28.7 (±0.1) cm TL and 506.6 (±7.1) g, respectively. The slope b of the length-weight relationship was 2.98, 3.01 3.01, for males, females, and combined sexes, respectively. The relative condition factor was 1.02 for males and 1.04 for females with little variation across the months of sampling. The length at 50% maturity was estimated as 31.0 cm TL for male Nile tilapia and 26.0 cm TL for females. Sixty percent of the fish in the commercial catches surveyed were below 30 cm TL. Comparisons with earlier studies in this system suggest an overall decline in size at maturity over the past 30 years, which may reflect intense fishing pressure.     

Feeding ecology of a bagrid catfish,Mystus tengara (Hamilton, 1822) in the Tanore wetland of Rajshahi,Northwestern Bangladesh

The feeding ecology of Mystus tengara (Hamilton, 1822) was studied in the Tanore wetland of Rajshahi, northwestern Bangladesh from January 2011 to December 2011. In fish purchased from the local market, a total of 14 713 food items were identified in the stomach contents of 3191 M. tengara specimens, an omnivorous catfish with a diversified feeding strategy including cladocerans, copepods, rotifers, gastropods, green algae, diatoms, insects, ‘small fish remains, fish fry and eggs’ and ‘other’ food items. Insects were the most dominant group of prey throughout the year. Seasonal changes in diet composition were due to the availability of prey items and fish size. Ontogenetic changes were noted between the juveniles and adults. The gut contents of M. tengara showed that the food items were from different layers of the water bodies.     

Diet overlap of non‐native common carp and native Nile tilapia: A potential cause for population reduction of Nile tilapia stock in Lake Ziway,Ethiopia

Lake Ziway harbours indigenous and exotic fish species including Nile tilapia (Oreochromis niloticus) and common carp (Cyprinus carpio). Nile tilapia was the dominant and preferred fish. However, its contribution to total catch has dramatically declined from 89.3% in 1994 to 27% in 2014 while the introduced common carp has increased from 0% before 2012 to 25% in 2014. Common carp potentially compete with Nile tilapia for available resources and could be a cause for the decline. Thus, the study explored the dietary overlap of the two species from April to August 2017. Schoener's overlap index (α) revealed significant dietary overlap between the two species (α = 0.84, between juveniles and α = 0.63, between adults). Juveniles fed mainly on animal origin (zooplankton and insects), while adult Nile tilapia have consumed plant origin (macrophyte and phytoplankton), and adult common carp fed on macrophytes and detritus. The presence of significant dietary overlap between the two species, particularly due to intense competition among juveniles, might cause the reduction of Nile tilapia stock because the native species has shown a competitive disadvantage for food in the presence of common carp. This study provides baseline information to researchers and decision makers working towards the sustainable resource utilisation of the system.     

Effect of dietary fish meal replacement by red algae, Gracilaria arcuata, on growth performance and body composition of Nile tilapia Oreochromis niloticus

A 12-week long feeding experiment was initiated to evaluate the effect of dietary supplementation of red algae, Gracilaria arcuata, on the growth performance, feed utilization and body composition of Nile tilapia Oreochromis niloticus (Linnaeus, 1758). The fish were fed with an algae-free control diet (C) and three experimental diets which replaced conventional fish meal with varying levels of dried G. arcuata (20%, 40% and 60%, represented as G20, G40 and G60, respectively). The growth parameters of final weight (FW), weight gain (WG), percentage of weight gain (WG%), daily growth rate (DGR) and specific growth rate (SGR) were significantly reduced (P < 0.05) at all levels of algae incorporation compared to the control diet. Moreover, the negative impact of Gracilaria meal on the growth performance of Nile tilapia increased as the proportion of algae in the diet increased, with fish on diet G20 exhibiting a significantly higher growth performance than the fish on either of the G40 and G60 diets. On the other hand, the feed utilization parameters feed conversion ratio (FCR) and protein efficiency ratio (PER) did not show significant differences between the fish in the control group and those on diet G20, although poorer FCR and PER outcomes were achieved in the case of fish on diet G60. The content of moisture, protein and ash in muscle and carcass increased as the proportion of Gracilaria meal in the diets increased, but the reverse was true for lipid level. These results indicate that incorporation of less than 20% red algae, Gracilaria arcuata, could be feasible in the diet of Nile tilapia and further studies are recommended to optimize the level of algae to improve growth performance.     

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.     

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     

Effect of water temperature,feeding frequency,and protein percent in the diet on water quality,growth and behavior of Nile tilapia Oreochromis niloticus (Linnaeus, 1758)

This study evaluated how water temperature (26, 28, and 30°C), number of meals per day (one or two meals), and protein percent in diet (20, 25 and 30%) impact growth performance, biometric indices, and feeding behavior of Nile tilapia, Oreochromis niloticus. Fish were randomly allocated into 18 equal replicate groups. Higher final body weight was observed in fish reared at 30°C and fed one meal per day containing 30% crude protein. Better weight gain, weight gain %, feed conversion ratio, specific growth rate, and condition factor were recorded in fish reared at 26°C and fed one meal per day containing 30% protein. The best length weight relationship was obtained in fish reared at 26°C and fed one meal per day containing 30% crude protein. Shorter feeding duration and duration of appetite inhibition latency were recorded in fish reared at 30°C, fed one meal per day, and given a diet containing 30% protein. The highest proactivity was recorded in fish reared at 30°C, received one meal per day, and with 25% crude protein in their diet. Conclusively, rearing Nile tilapia at 26–30°C with a lower feeding frequency (one meal/day) and a 30% crude protein diet achieved better performance and feeding behavior.     

Food and feeding habits of Clarias mossambicus from four areas in the Lake Victoria basin,East Africa

Synopsis The food and feeding habits of Clarias mossambicus in four habitat types in the northern Lake Victoria basin were studied. The importance of certain food types to this species varies from area to area. In the lake, rivers and swamps, C. mossambicus fed mainly on fish, particularly Haplochromis species. In the sewage ponds, Crustacea and insects were the main food items. The extent of ingestion by C. mossambicus of a given food type presumably reflects the availability and abundance of the food in the habitat. This constitutes a shift in the food habits which is of vital importance in this migratory fish. By employing several feeding methods, i.e. hunting, bottom feeding, surface feeding and filter feeding, C. mossambicus exploits a broad spectrum of food resources in the various habitats.Many modern workers consider Clarias mossambicus conspecific with C. gariepinus and a junior synonym of the latter (Editor)     

The introduced Oreochromis niloticus is spreading on the Kaufe floodplain,Zambia

The Nile tilapia, Oreochromis niloticus, escaped from a fish farm in Mazabuka, probably in 1987, and was found in September 1992 on a 75 km stretch of the river, mainly downstream from the site of release. In most areas, except near Mazabuka, O. niloticus was relatively uncommon and so far does not seem to have hybridized with the local tilapias.