Trophic complexity of small fish in nearshore food webs

Small nearshore fishes are an important part of lacustrine and functional diversity and link pelagic and benthic habitats by serving as prey for larger nearshore and offshore fishes. However, the trophic complexity of these small nearshore fishes is often unrecognized and detailed studies of their role in food webs are lacking. Here, we examined niche space patterns of small nearshore fish species using Bayesian analyses of carbon and nitrogen stable isotope data in nine freshwater lakes that are among the largest lakes in Minnesota. We found considerable variability in niche areas within species and high variability in niche overlap across species. At the assemblage level, niche overlap (average diet overlap of all species pairs at a lake) decreased as whole-lake species richness increased, possibly indicating a greater degree of resource specialization in more speciose lakes. Overall fish niche space was weakly but significantly related to niche space of their invertebrate prey. Although nearshore benthic resources contributed to fish diets in all lakes, all fish species also had non-negligible and variable contributions from pelagic zooplankton. This inter- and intraspecific variability in trophic niche space likely contributes to the multi-level trophic complexity, functional diversity, and potentially food web resilience to ecosystem changes.

     

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.     

Small pelagic fish assemblages in relation to environmental regimes in the Central Mediterranean

Stable isotope analyses and derived population-level metrics were used to quantitatively analyse spatial and seasonal heterogeneity in the fish trophic dynamics in relation to environmental variables in Mwanza Gulf, Lake Victoria (Tanzania). The fish community in Lake Victoria, including the top predator Nile perch, is generally omnivorous with a heavy reliance on invertebrates. This is in contrast to findings based on stomach content analyses of Nile perch, which showed a stronger reliance on fish. We tested two hypotheses: (1) during the rainy seasons multiple carbon sources influence the food-web structure inside the Gulf, leading to increased carbon ranges and trophic diversity. (2) During dry periods, the food-web structure mainly relies on pelagic primary production, reducing carbon ranges and trophic diversity. Carbon sources indeed varied seasonally and spatially, affecting the fish community at the highest trophic levels. With the onset of rains, carbon sources became spatially highly differentiated with enriched δ¹³C values of fish in shallow water inside the Gulf and depleted δ¹³C values in open waters. Metrics associated with niche size correlated significantly with seasonally varying environmental variables, while δ¹³C ranges correlated with spatially varying environmental variables.     

Changes in the feeding biology of the Nile perch, Lates niloticus (L.) (Pisces: Centropomidae), in Lake Victoria, East Africa since its introduction in 1960, and its impact on the native fish community of the Nyanza Gulf

Lates niloticus is not native to Lake Victoria but was introduced during or shortly before 1960. It remained relatively uncommon until 1975, when the number in the Nyanza Gulf began to increase impressively, the estimated catch rising over 100-fold between 1978 and 1982. Originally Lates was piscivorous, its diet reflecting the composition of the native fish community. The present investigation has revealed that its diet is now almost entirely comprised of Caridina nilotica , a small microphagous prawn, and juvenile Lates. Native fish species, except for the small pelagic Rastrineobola argenteus , are very rarely consumed. This change in diet is a result of the shattering impact Lates predation has had on the native fishes, which have been virtually wiped out. The original community, which was dominated by several hundred haplochromine species and the catfishes Clarias mossambicus and Bagrus docmac which preyed upon them, and included two endemic tilapiine cichlids and 38 species of non-cichlids, no longer exists. It has been replaced by a community dominated by Lates which now accounts for well over 80% of the fish biomass in the Nyanza Gulf and very nearly 100% in the study area. The only other species regularly encountered were Oreochromis niloticus , an introduced tilapiine, and Rastrineobola argenteus , a native zooplanktivore.     

Diet composition,feeding niche partitioning and trophic organisation of large pelagic predatory fishes in the eastern Arabian Sea

Information on the ecology and feeding behaviour of the large oceanic predatory fishes is crucial for the ecosystem approaches to fisheries management models. Co-existing large pelagic predators in the open oceans may avoid competition for the limited forage by resource partitioning on spatial, temporal or trophic levels. To test this, we studied the prey species composition, diet overlap, trophic level, and trophic organisation of 12 large predatory fishes co-existing in the eastern Arabian Sea. Stomach contents of 1,518 specimens caught by exploratory longline operations in the Indian Exclusive Economic Zone during the years 2006–2009 were analysed. Finfishes were dominant prey of all species except blue marlin (Makaira nigricans) and yellowfin tuna (Thunnus albacares), which fed mainly on cephalopods, and long-snouted lancetfish (Alepisaurus ferox) and pelagic stingray (Pteroplatytrygon violacea), which fed mainly on crustaceans. Common dolphinfish (Coryphaena hippurus) and yellowfin tuna fed on a wider variety of prey than the other species, while the diets of lancetfish and black marlin (Istiompax indica) were narrowest. Pelagic stingray and great barracuda (Sphyraena barracuda) fed on species occupying epipelagic waters, whereas the contribution of mesopelagic prey was higher in the diets of swordfish (Xiphias gladius) and pelagic thresher (Alopias pelagicus). Trophic levels of these fishes ranged from 4.13 to 4.37. Diet overlap index revealed that some of the large pelagic predatory fishes share common prey species. Cluster analysis of the diets revealed four distinct trophic guilds namely ‘flyingfish feeders’ (common dolphinfish and great barracuda); ‘mesopelagic predators’ (pelagic thresher and swordfish); ‘crab feeders’ (lancetfish, pelagic stingray and silky shark) and ‘squid feeders’ (yellowfin tuna, Indo-Pacific sailfish (Istiophorus platypterus), skipjack tuna (Katsuwonus pelamis), black marlin and blue marlin). Large predatory fishes of the eastern Arabian Sea target different prey types, and limit their vertical extent and time of feeding to avoid competing for prey.     

Dietary niche partitioning in sympatric gadid species in coastal Newfoundland: evidence from stomachs and C-N isotopes

The feeding habits of co-occurring gadid species Atlantic cod (Gadus morhua) and Greenland cod (Gadus ogac) in coastal Newfoundland waters, examined using stable isotope (δ ¹³C and δ ¹⁵N) and stomach content analysis, indicated little dietary niche overlap and interspecific competition for food resources despite similar trophic levels. Both species consumed a variety of invertebrates and fish but showed a preference for different prey items. Polychaetes, fish and small crustaceans dominated G. ogac stomach contents while small crustaceans, in particular hyperiid amphipods and fish, dominated those of G. morhua. In general, G. morhua consumed more pelagic prey and had a significantly more pelagic (more negative) δ ¹³C signature while G. ogac consumed primarily benthic prey and had a more benthic (more positive) δ ¹³C signature. δ ¹⁵N levels were similar in these species suggesting similar trophic positions, with levels increasing with fish length in both species. Dietary overlap was not significant in both stomach and stable isotope analyses. We conclude that interspecific competition for food is low between G. ogac and G. morhua and is unlikely to be a factor in the slow rebuilding of Atlantic cod in this region.     

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.

     

Stable isotope evidence from formalin–ethanol-preserved specimens indicates dietary shifts and increasing diet overlap in Lake Victoria cichlids

Stable isotopes are increasingly being used to infer past and present trophic interactions in light of environmental changes. The Lake Victoria haplochromine cichlids have experienced severe environmental changes in the past decades that, amongst others, resulted in a dietary shift towards larger prey. We investigated how the changed environment and diet of the haplochromines influenced stable isotope values of formalin-then-ethanol-preserved cichlid specimens, and then investigated how these values differed among species before (1977–1982) and after substantial environmental changes (2005–2007). We found a small preservation effect on both δ¹³C and δ¹⁵N values, and significant differences in isotope values among haplochromine species collected before the environmental changes. In contrast, there was a remarkable similarity in δ¹³C and δ¹⁵N values among species collected from the contemporary ecosystem and two out three species showed significantly different stable isotope values compared to species of the historic ecosystem. In addition, we found a putative isotopic gradient effect along our 5-km-long research transect indicating that the studied demersal species are more stenotopic than previously thought. The environmental changes have resulted in dietary change and overlap of the haplochromines which provides insight into the trophic plasticity of these species, which are often considered trophic specialists.     

Trophic divergence of Lake Kivu cichlid fishes along a pelagic versus littoral habitat axis

Local adaptation to the littoral and pelagic zones in two cichlid haplochromine fish species from Lake Kivu was investigated using morphometrics. Cranial variation and inferred jaw mechanics in both sexes of the two species across the two habitat types were quantified and compared. Comparisons of littoral versus pelagic populations revealed habitat‐specific differences in the shape of the feeding apparatus. Also, kinematic transmission of the anterior jaw four‐bar linkage that promotes greater jaw protrusion was higher in the pelagic zone than in the littoral zone for both species. Inferred bite force was likewise higher in pelagic zone fish. There were also sex‐specific differences in craniofacial morphology as males exhibited longer heads than females in both habitats. As has been described for other cichlids in the East African Great Lakes, local adaptation to trophic resources in the littoral and pelagic habitats characterizes these two Lake Kivu cichlids. Similar studies involving other types of the Lake Kivu fishes are recommended to test the evidence of the observed trophic patterns and their genetic basis of divergences.     

Trophic structure of a coastal fish community determined with diet and stable isotope analyses

A combination of dietary guild analysis and nitrogen (δ¹⁵N) and carbon (δ¹³C) stable‐isotope analysis was used to assess the trophic structure of the fish community in Rhode Island and Block Island Sounds, an area off southern New England identified for offshore wind energy development. In the autumn of 2009, 2010 and 2011, stomach and tissue samples were taken from 20 fish and invertebrate species for analysis of diet composition and δ¹⁵N and δ¹³C signatures. The food chain in Rhode Island and Block Island Sounds comprises approximately four trophic levels within which the fish community is divided into distinct dietary guilds, including planktivores, benthivores, crustacivores and piscivores. Within these guilds, inter‐species isotopic and dietary overlap is high, suggesting that resource partitioning or competitive interactions play a major role in structuring the fish community. Carbon isotopes indicate that most fishes are supported by pelagic phytoplankton, although there is evidence that benthic production also plays a role, particularly for obligate benthivores such as skates Leucoraja spp. This type of analysis is useful for developing an ecosystem‐based approach to management, as it identifies species that act as direct links to basal resources as well as species groups that share trophic roles.     

Food‐web structure and mercury dynamics in a large subarctic lake following multiple species introductions

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Depth gradients in food‐web processes linking habitats in large lakes: Lake Superior as an exemplar ecosystem

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昆承湖优势种鱼类时空-营养生态位

为了解昆承湖优势种鱼类资源利用情况,首先利用生态位方法计算了时间、空间及营养三个资源维度的生态位宽度及重叠值,然后根据时空-营养生态位宽度值将优势种鱼类划分为广位种、中位种和窄位种,最后讨论了生态位宽度及重叠的可能原因。结果显示：刀鲚Coilia nasus、蒙古鲌Chanodichthys mongolicus、似鱎Toxabramis swinhonis、似鳊Pseudobrama simoni、鳙Hypophthalmichthys nobilis、鲢Hypophthalmichthys molitrix、花鱼骨Hemibarbus maculatus、似刺鳊鮈Paracanthobrama guichenoti、大鳍鱊Acheilognathus macropterus和鲫Carassius auratus为优势种。在时间维度：鲢的生态位宽度最大,似鳊的最小;生态位重叠具有显著意义的有24对,占总的53.33%。在空间维度,似刺鳊鮈最大,鲢最小;生态位重叠具有显著意义的有36对,占总的80%。在营养维度,最大的为鲫,最小的为花鱼骨;生态位重叠具有显著意义的有8对,占17.78%...     

Wasp-waist interactions in the North Sea ecosystem

Background

In a “wasp-waist” ecosystem, an intermediate trophic level is expected to control the abundance of predators through a bottom-up interaction and the abundance of prey through a top-down interaction. Previous studies suggest that the North Sea is mainly governed by bottom-up interactions driven by climate perturbations. However, few studies have investigated the importance of the intermediate trophic level occupied by small pelagic fishes.

Methodology/Principal Findings

We investigated the numeric interactions among 10 species of seabirds, two species of pelagic fish and four groups of zooplankton in the North Sea using decadal-scale databases. Linear models were used to relate the time series of zooplankton and seabirds to the time series of pelagic fish. Seabirds were positively related to herring (Clupea harengus), suggesting a bottom-up interaction. Two groups of zooplankton; Calanus helgolandicus and krill were negatively related to sprat (Sprattus sprattus) and herring respectively, suggesting top-down interactions. In addition, we found positive relationships among the zooplankton groups. Para/pseudocalanus was positively related to C. helgolandicus and C. finmarchicus was positively related to krill.

Conclusion/Significance

Our results indicate that herring was important in regulating the abundance of seabirds through a bottom-up interaction and that herring and sprat were important in regulating zooplankton through top-down interactions. We suggest that the positive relationships among zooplankton groups were due to selective foraging and switching in the two clupeid fishes. Our results suggest that “wasp-waist” interactions might be more important in the North Sea than previously anticipated. Fluctuations in the populations of pelagic fish due to harvesting and depletion of their predators might accordingly have profound consequences for ecosystem dynamics through trophic cascades.     

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.     

Discrete foraging niches promote ecological, phenotypic, and genetic divergence in sympatric whitefish (Coregonus lavaretus)

Natural populations often vary in their degree of ecological, morphological and genetic divergence. This variation can be arranged along an ecological speciation continuum of increasingly discrete variation, with high inter-individual variation at one end and well defined species in the other. In postglacial fishes, evolutionary divergence has commonly resulted in the co-occurrence of a pelagic and a benthic specialist. We studied three replicate lakes supporting sympatric pelagic and benthic European whitefish (Coregonus lavaretus (L.)) morphs in search for early signs of possible further divergence into more specialized niches. Using stomach content data (recent diet) and stable isotope analyses (time-integrated measure of trophic niche use), we observed a split in the trophic niche within the benthic whitefish morph, with individuals specializing on either littoral or profundal resources. This divergence in resource use was accompanied by small but significant differences in an adaptive morphological trait (gill raker number) and significant genetic differences between fish exploiting littoral and profundal habitats and foraging resources. The same pattern of parallel divergence was found in all three lakes, suggesting similar natural selection pressures driving and/or maintaining the divergence. The two levels of divergence (a clear and robust benthic – pelagic and a more subtle littoral – profundal divergence) observed in this study apparently represent different stages in the process of ecological speciation.     

Trophic plasticity in the Atlantic sharpnose shark (Rhizoprionodon terraenovae) from the north central Gulf of Mexico

Quantifying the trophic role of sharks in coastal ecosystems is crucial for the construction of accurate ecosystem models. This is particularly important for wide-ranging species like the Atlantic sharpnose shark (Rhizoprionodon terraenovae), ubiquitous across the northern Gulf of Mexico. We used gut content and stable isotope analyses to determine if differences in abundance of Atlantic sharpnose sharks in the waters around Mobile Bay, Alabama translated into differences in dietary sources or trophic position among sharks sampled east and west relative to the mouth of the bay. Gut content analysis suggested that Atlantic sharpnose sharks eat primarily teleost fishes (%IRI?>?90% across size classes), and both stomach content and stable isotope analyses highlighted an ontogenetic shift in diet. Nitrogen stable isotope data from liver and muscle tissues indicated regional shifts in trophic position for Atlantic sharpnose sharks. The mixing model SIAR (stable isotope analysis in R) v.4.0.2 was used to suggest possible contributions from likely prey items for Atlantic sharpnose sharks sampled east and west of Mobile Bay. Portunid crabs and shrimp made higher contributions to the diet of Atlantic sharpnose sharks in the western region, compared to higher and more variable contributions from fish like croaker (Micropogonias undulatus) and hardhead catfish (Arius felis) in the eastern region. Our results suggest trophic plasticity in Atlantic sharpnose sharks, findings that emphasize the importance of examining regional variation in trophic position when constructing coastal foodweb models.     

Food web overlap among native axolotl (<Emphasis Type="Italic">Ambystoma mexicanum</Emphasis>) and two exotic fishes: carp (<Emphasis Type="Italic">Cyprinus carpio</Emphasis>) and tilapia (<Emphasis Type="Italic">Oreochromis niloticus</Emphasis>) in Xochimilco,Mexico City

Two exotic fishes, common carp (Cyprinus carpio) and tilapia (Oreochromis niloticus), were introduced more than 20 years ago into Xochimilco for aquaculture, and now dominate the system in terms of biomass and numbers. Over this same period, wild populations of the microendemic axolotl salamander (Ambystoma mexicanum) have been dramatically reduced, and it currently persists in isolated areas of this aquatic system, which is situated at the edge of Mexico City. In this study, we examine potential trophic interactions and niche overlap among two exotic fishes: carp and tilapia, and the native axolotl. Axolotl had more diverse diets and a higher trophic position compared to the exotics. Stable isotope analysis revealed substantial trophic niche overlap among axolotl and the exotics. The two exotics occupied a larger niche area than the axolotl, suggesting higher levels of omnivory and diet specialization. Current exotic fish removal efforts will further our understanding of interactions between the axolotl and exotic species, and are expected to benefit axolotl recovery efforts.     

Co-introduction of Dolicirroplectanum lacustre,a monogenean gill parasite of the invasive Nile perch Lates niloticus: intraspecific diversification and mitonuclear discordance in native versus introduced areas

The Nile perch (Lates niloticus) is a notorious invasive species. The introductions of Nile perch into several lakes and rivers in the Lake Victoria region led to the impoverishment of trophic food webs, particularly well documented in Lake Victoria. Additionally, its parasites were co-introduced, including Dolicirroplectanum lacustre (Monogenea, Diplectanidae). Dolicirroplectanum lacustre is the single monogenean gill parasite of latid fishes (Lates spp.) inhabiting several major African freshwater systems. We examined the intra-specific diversification of D. lacustre from Lates niloticus in Lake Albert, Uganda (native range) and Lake Victoria (introduced range) by assessing morphological and genetic differentiation, and microhabitat preference. We expected reduced morphological and genetic diversity for D. lacustre in Lake Victoria compared with Lake Albert, as a result of the historical introductions. We found that D. lacustre displayed high morphological variability within and between African freshwaters, with two morphotypes identified, as in former studies. The single shared morphotype between Lake Albert and Lake Victoria displayed similar levels of haplotype and nucleotide diversity between the lakes. Mitonuclear discordance within the morphotypes of D. lacustre indicates an incomplete reproductive barrier between the morphotypes. The diversification in the mitochondrial gene portion is directly linked with the morphotypes, while the nuclear gene portions indicate conspecificity. Based on our results, we reported reduced genetic and morphological diversity, potentially being a result of a founder effect in Lake Victoria.     

High prevalence of non-synonymous substitutions in mtDNA of cichlid fishes from Lake Victoria

When a population size is reduced, genetic drift may fix slightly deleterious mutations, and an increase in nonsynonymous substitution is expected. It has been suggested that past aridity has seriously affected and decreased the populations of cichlid fishes in Lake Victoria, while geographical studies have shown that the water levels in Lake Tanganyika and Lake Malawi have remained fairly constant. The comparably stable environments in the latter two lakes might have kept the populations of cichlid fishes large enough to remove slightly deleterious mutations. The difference in the stability of cichlid fish population sizes between Lake Victoria and the Lakes Tanganyika and Malawi is expected to have caused differences in the nonsynonymous/synonymous ratio, ω (= d_N/d_S), of the evolutionary rate. Here, we estimated ω and compared it between the cichlids of the three lakes for 13 mitochondrial protein-coding genes using maximum likelihood methods. We found that the lineages of the cichlids in Lake Victoria had a significantly higher ω for several mitochondrial loci. Moreover, positive selection was indicated for several codons in the mtDNA of the Lake Victoria cichlid lineage. Our results indicate that both adaptive and slightly deleterious molecular evolution has taken place in the Lake Victoria cichlids' mtDNA genes, whose nonsynonymous sites are generally conserved.