Stable isotopes indicate that zebra mussels (Dreissena polymorpha) increase dependence of lake food webs on littoral energy sources

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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 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 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.     

Isotopic evidence of functional overlap amongst the resilient pelagic fishes

The Lake Victoria ecosystem once hosted a diverse fish community dominated by a large species flock of haplochromine cichlids. Today this fish assemblage is highly altered by anthropogenic activities, with at least half of the indigenous species either extinct or very rare. The fauna and flora of the tropic’s largest lake are still in flux, and little is known about even the most basic ecological questions, such as the source of carbon and nitrogen for its spectacular productivity. The actual food-web structure is difficult to determine with results based on traditional gut analysis. At the moment both scientists and conservationists are at odds as to whether there is any evidence of functional replacement or functional overlap amongst the resilient pelagic fishes of Lake Victoria. This study used a static stable isotope model to investigate the source of carbon and the extent of the trophic overlap amongst three pelagic fishes, Rastrineobola argentea, ssichromis laparogramma and Y. fusiformis in the Kenyan waters of Lake Victoria. The δ ¹³C of plants ranged from ?8.8 to ?24.6 $\permil$, while fishes ranged from ?18.6 to ?24.5 $\permil$, suggesting assimilation of mostly C₃ sources for the fish species. On the basis of the isotope model, it is evident that Yssichromis laparogramma, Yssichromis fusiformis , and Rastrineobola argentea obtained from 90 to 97.5$%$, with an average of 94.2$%$, of their food from the same trophic level. The very high degree of overlap in the effective trophic level of these three zooplanktivores has important ecological and conservation implications.     

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.     

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.     

Role of body size and temporal hydrology in the dietary shifts of shortjaw tapertail anchovy Coilia brachygnathus (Actinopterygii, Engraulidae) in a large floodplain lake

Seasonal water-level changes in floodplain lakes can induce variations in primary and secondary production, thus affecting trophic interactions. In this study, we tested the latter by studying size- and temporal hydrology-related shifts in the diet of shortjaw tapertail anchovy Coilia brachygnathus (Actinopterygii, Engraulidae) from Lake Poyang. During the wet season, δ¹³C values ranged from ?28.2‰ for small anchovies to ?24.6‰ for larger individuals, but δ¹⁵N ranged from 18.9‰ for smaller fish to 12.4‰ for larger fish. Significant ¹³C-enrichment and ¹⁵N-depletion occurred with increasing size, revealing that different carbon sources were used as the fish grew. Given the high hydrologic fluctuation levels, significant differences in δ¹³C values were observed among larger anchovies between seasons, indicating a temporal dietary shift. Anchovies fed primarily on shrimp and fish during the low-water season despite the predominance of zooplankton during the two seasons studied, which indicated increased piscivorous reliance. C. brachygnathus exhibited higher δ¹⁵N values during the wet season because the food items were ¹⁵N-enriched. Human waste brought by floods could be another possible interpretation. Considering C. brachygnathus is an important link between plankton production and higher piscivorous trophic levels, changes in the species are expected to affect the functioning of lake food webs along the trophic pathway.     

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.     

Trophic interactions in the Zoige Alpine wetland on the eastern edge of the Qinghai–Tibetan Plateau inferred by stable isotopes

The Zoige wetland ecosystem of the eastern edge of the Qinghai–Tibetan Plateau is the largest plateau peat bog worldwide and an important biodiversity conservation centre in China. Nevertheless, limited information is available about this Alpine wetland aquatic ecosystem, including its food-web structure and complexity. In this study, we analysed the trophic interactions among dominant organisms in the food web of this Alpine wetland aquatic ecosystem and assessed the importance of allochthonous and autochthonous carbon sources during the wet season using δ¹³C and δ¹⁵N analysis. The results supported the hypothesis that the food web of the Zoige Alpine wetland is largely maintained by allochthonous carbon sources rather than autochthonous food sources during the wet season when the allochthonous resource availability is relatively high. The food web contains approximately three trophic levels, and Schizopygopsis pylzovi (a genus of cyprinid fish) is the top predator. Intermediate trophic positions, including zooplankton and macroinvertebrates, represent the primary animal community in the food web. The allochthonous carbon sources are transferred to the food web via these macroinvertebrates (e.g., Ecdyonurus sp. and Sigara substriata). Although our findings show a low biodiversity in the Zoige wetland during the wet season, the trophic redundancy in the food-web structure could support the stability of the Zoige Alpine wetland aquatic ecosystem.     

Long‐term impacts of invasive species on a native top predator in a large lake system

1. Declining abundances of forage fish and the introduction and establishment of non‐indigenous species have the potential to substantially alter resource and habitat exploitation by top predators in large lakes. 2. We measured stable isotopes of carbon (δ¹³C) and nitrogen (δ¹⁵N) in field‐collected and archived samples of Lake Ontario lake trout (Salvelinus namaycush) and five species of prey fish and compared current trophic relationships of this top predator with historical samples. 3. Relationships between δ¹⁵N and lake trout age were temporally consistent throughout Lake Ontario and confirmed the role of lake trout as a top predator in this food web. However, δ¹³C values for age classes of lake trout collected in 2008 ranged from 1.0 to 3.9‰ higher than those reported for the population sampled in 1992. 4. Isotope mixing models predicted that these changes in resource assimilation were owing to the replacement of rainbow smelt (Osmerus mordax) by round goby (Neogobius melanostomus) in lake trout diet and increased reliance on carbon resources derived from nearshore production. This contrasts with the historical situation in Lake Ontario where δ¹³C values of the lake trout population were dominated by a reliance on offshore carbon production. 5. These results indicate a reduced capacity of the Lake Ontario offshore food web to support the energetic requirements of lake trout and that this top predator has become increasingly reliant on prey resources that are derived from nearshore carbon pathways.     

Did the loss of phytoplanktivorous fish contribute to algal blooms in the Mwanza Gulf of Lake Victoria?

Possible causes of the increased algal blooms in Lake Victoria in the 1980s have been disputed by several authors; some suggested a top-down effect by the introduced Nile perch, whereas others suggested a bottom-up effect due to eutrophication. In this article the potential impact is established of grazing by fish on phytoplankton densities, before the Nile perch upsurge and the concomitant algal blooms in the Mwanza Gulf. The biomass and trophic composition of fish in the sublittoral area of the Mwanza Gulf were calculated based on catch data from bottom trawls, and from gill nets covering the whole water column. Estimates of phytoplankton production in the same area were made from Secchi values and chlorophyll concentrations. The total phytoplankton intake by fish was estimated at 230 mg DW m⁻² day⁻¹. The daily gross production ranged from 6,200 to 7,100 mg DW m⁻² day⁻¹ and the net production from 1,900 to 2,200 mg DW m⁻² day⁻¹. Thus, losses of phytoplankton through grazing by fish were about 3–4% of daily gross and 10–12% of daily net phytoplankton production. As a consequence it is unlikely that the phytoplankton blooms in the second half of the 1980s were due to a top-down effect caused by a strong decline in phytoplankton grazing by fish.     

Impacts of zebra mussels (Dreissena polymorpha) on isotopic niche size and niche overlap among fish species in a mesotrophic lake

Zebra mussels (Dreissena polymorpha) filter feed phytoplankton and reduce available pelagic energy, potentially driving fish to use littoral energy sources in lakes. However, changes in food webs and energy flow in complex fish communities after zebra mussel establishment are poorly known. We assessed impacts of zebra mussels on fish littoral carbon use, trophic position, isotopic niche size, and isotopic niche overlap among individual fish species using δ¹³C and δ¹⁵N data collected before (2014) and after (2019) zebra mussel establishment in Lake Ida, MN. Isotope data were collected from 11 fish species, and from zooplankton and littoral invertebrates to estimate baseline isotope values. Mixing models were used to convert fish δ¹³C and δ¹⁵N into estimates of littoral carbon and trophic position, respectively. We tested whether trophic position, littoral carbon use, isotopic niche size, and isotopic niche overlap changed from 2014 to 2019 for each fish species. We found few effects on fish trophic position, but 10 out of 11 fish species increased littoral carbon use after zebra mussel establishment, with mean littoral carbon increasing from 43% before to 67% after establishment. Average isotopic niche size of individual species increased significantly (2.1-fold) post zebra mussels, and pairwise-niche overlap between species increased significantly (1.2-fold). These results indicate zebra mussels increase littoral energy dependence in the fish community, resulting in larger individual isotopic niches and increased isotopic niche overlap. These effects may increase interspecific competition among fish species and could ultimately result in reduced abundance of species less able to utilize littoral energy sources.

     

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.     

Lake Victoria fisheries: the Kenyan reality and environmental implications

Synopsis The introduction of the Nile perch into Lake Victoria has dramatically altered the fishery in that lake and contributed to the decline of the fishery for indigenous tilapias. One sector of the fishery in Lake Victoria has benefitted from the Nile perch introduction, although catches have declined in recent years. Inefficient enforcement of fisheries regulations has had a detrimental effect on indigenous species but may also have contributed to the recent decline in Nile perch catches. Fisheries development plans have tended to favour capital-intensive fisheries and to ignore small scale subsistence fisheries. A case study at Wichlum Beach on the Kenyan shores of Lake Victoria has revealed the efficiency of traditional fishing and fish drying methods as well as the high ecological costs of the practice of kiln-drying Nile perch. Forty-five tons of firewood are used per month at Wichlum Beach alone for kiln-drying perch. The increased economic viability of the fishery has attracted professionals into the industry and resulted in the development of an export-oriented trade. The Yala Swamp adjacent to Lake Victoria has been extensively drained as part of a large land reclamation scheme and more draining is planned. Increased environmental awareness in Kenya, and Kenya's membership in the Convention on Wetlands of International Importance, has resulted in a critical review of these plans.     

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.     

Divergence in aerobic scope and thermal tolerance is related to local thermal regime in two populations of introduced Nile perch (Lates niloticus)

We tested whether thermal tolerance and aerobic performance differed between two populations of Nile perch (Lates niloticus) originating from the same source population six decades after their introduction into two lakes in the Lake Victoria basin in East Africa. We used short-term acclimation of juvenile fish to a range of temperatures from ambient to +6°C, and performed critical thermal maximum (CT_max) and respirometry tests to measure upper thermal tolerance, resting and maximum metabolic rates, and aerobic scope (AS). Across acclimation temperatures, Nile perch from the cooler lake (Lake Nabugabo, Uganda) tended to have lower thermal tolerance (i.e., CT_max) and lower aerobic performance (i.e., AS) than Nile perch from the warmer waters of Lake Victoria (Bugonga region, Uganda). Effects of temperature acclimation were more pronounced in the Lake Victoria population, with the Lake Nabugabo fish showing less thermal plasticity in most metabolic traits. Our results suggest phenotypic divergence in thermal tolerance between these two introduced populations in a direction consistent with an adaptive response to local thermal regimes.     

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.     

Fifty years later: trophic ecology and niche overlap of a native and non-indigenous fish species in the western basin of Lake Erie

Since the introduction of white perch (Morone americana) into Lake Erie over 50 years ago, the population size of native yellow perch (Perca flavescens) has decreased up to 79 % and significant changes to the ecosystem have occurred. We examined long-term population estimates and used stable isotopes of carbon (δ ¹³C) and nitrogen (δ ¹⁵N) paired with stomach content analysis to quantify the trophic ecology and niche overlap of adult yellow perch and white perch in the western basin of Lake Erie. We found that changes in yellow perch abundance since 1979 appeared to be better correlated with changes in fishery exploitation rates than with food competition effects from white perch. At the time of this study, yellow perch were found to have higher δ ¹³C values, indicating greater utilization of benthic food resources than white perch, and white perch occupied higher trophic positions based on δ ¹⁵N. The diets of both species varied spatially and seasonally based on stable isotopes and stomach contents, likely driven by changes in prey abundance. Comparison of niche widths using stable isotope population metrics and Schoener diet similarity index suggested a low to moderate degree of niche overlap between species. Isotopic niches of white perch were generally larger than those of yellow perch demonstrating broader resource utilization by this non-indigenous species. We submit that isotopic niche overlap comparisons are more appropriate for studies seeking to understand interactions among populations over course temporal scales, while diet overlap indices, such as the Schoener index provide a means to study fine-scale interactions such as ontogenetic and seasonal diet shifts.