Reciprocal trophic niche shifts in native and invasive fish: salmonids and galaxiids in Patagonian lakes

1. Rainbow (Oncorhynchus mykiss) and brown trout (Salmo trutta) are widespread and invasive salmonids with important lethal effects as predators, although indirect effects are also possible. We used stable isotope analyses (δ¹⁵N, δ¹³C) to explore how the density of invasive trout in 25 Patagonian lakes alters the trophic niche (TN) of a widespread native fish, Galaxias platei (Galaxiidae). We also explored how the density of the galaxiid influences the TN of invasive trout. 2. We quantified two aspects of the TN: (i) the proportion of littoral carbon (PL) and (ii) trophic height (TH) (i.e. the ‘height’ at which the fish feeds in the food web). We related these measures of TN in a given species to the density of other species (as estimated by catch‐per‐unit‐effort). 3. As G. platei body size increased, their PL increased (increasing littoral feeding) in several lakes. However, none of the fish species investigated showed changes in PL with increasing density of the other fish species. TH increased with body size in all three species. In addition, the TH of large G. platei declined with increasing trout density and, reciprocally, the TH of large S. trutta decreased with decreasing G. platei density. 4. The reciprocal effects of native and the invasive fish on TH were as large as a shift of one trophic level. This pattern is consistent with an exhaustion of galaxiid prey for both piscivorous G. platei and S. trutta in lakes with high trout density. 5. These finding support the suggested management strategy of culling trout from overpopulated lakes, which should simultaneously protect native fish and enhance a lucrative sport fishery for large trout.     

Trophic interactions between native and introduced fish species in a littoral fish community

The trophic interactions between 15 native and two introduced fish species, silverside Odontesthes bonariensis and rainbow trout Oncorhynchus mykiss, collected in a major fishery area at Lake Titicaca were explored by integrating traditional ecological knowledge and stable‐isotope analyses (SIA). SIA suggested the existence of six trophic groups in this fish community based on δ¹³C and δ¹⁵N signatures. This was supported by ecological evidence illustrating marked spatial segregation between groups, but a similar trophic level for most of the native groups. Based on Bayesian ellipse analyses, niche overlap appeared to occur between small O. bonariensis (<90 mm) and benthopelagic native species (31·6%), and between the native pelagic killifish Orestias ispi and large O. bonariensis (39%) or O. mykiss (19·7%). In addition, Bayesian mixing models suggested that O. ispi and epipelagic species are likely to be the main prey items for the two introduced fish species. This study reveals a trophic link between native and introduced fish species, and demonstrates the utility of combining both SIA and traditional ecological knowledge to understand trophic relationships between fish species with similar feeding habits.     

Piscivory and diet overlap between two non‐native fishes in southern Chilean streams

Trophic relations among introduced species may induce highly variable and complex effects in communities and ecosystems. However, studies that identify the potential impacts for invaded systems and illuminate mechanisms of coexistence with native species are scarce. Here, we examined trophic relations between two introduced fishes in streams of NW Patagonia, rainbow trout (Oncorhynchus mykiss) and brown trout (Salmo trutta). These species originate from different regions of the Northern Hemisphere but they now coexist as invading species over the world. We used gastric contents and stable isotopes analysis to compare the diets of two size‐classes of these two invaders in three localities of southern Chile. Both species displayed similar ontogenic diet shifts with smaller trout consuming mostly invertebrates and larger trout being more piscivorous and epibenthic feeders. However, piscivory was more prevalent in brown trout than in rainbow trout and highest at the site with the greatest density of native fishes suggesting that the availability of native fishes as trout prey may limit the occurrence of trout piscivory. We found an elevated dietary overlap between the two trout species at larger sizes while at smaller size a higher intraspecific dietary overlap occurred suggesting a potential interference competition among the two fish invaders especially at larger sizes. Our results highlight that the impacts of invading species on non‐native fishes are context specific (i.e. species and ontogenic stages) and thus, difficult to generalize.     

Dietary ontogeny and niche shift to piscivory in lacustrine brown trout Salmo trutta revealed by stomach content and stable isotope analyses

The feeding ecology and ontogeny of a large size range of brown trout Salmo trutta in Lake Fyresvatnet, southern Norway, were examined by stomach content and stable isotope analyses. According to the stomach contents, the S. trutta changed their diet at c. 30 cm total length (L_T). The smaller size classes fed on benthic invertebrates and surface insects, whereas larger S. trutta (>30 cm) fed mainly on whitefish Coregonus lavaretus. A similar, but more gradual shift to piscivory in the size range 25–30 cm was found when using the stable isotope mixing model SIAR to reveal dietary ontogeny. The δ¹³C isotopic signature confirmed that S. trutta independent of size predominantly relied upon benthic energy sources, suggesting that the littoral zone was the primary foraging habitat for both invertebrate and piscivorous feeders. The δ¹⁵N values and trophic position increased with predator length, ranging from an average of 3·60 for small‐sized S. trutta (<15 cm) to 4·15 for large‐sized fish (>35 cm). The S. trutta exhibited a relatively slow growth rate during the predominant invertebrate feeding stages up to 7 years of age and 28 cm L_T, whereas fish above this size and age displayed a rapid growth rate of 9–11 cm year^?1, demonstrating the profitability of piscivorous feeding.     

Co‐varying impacts of land use and non‐native brown trout on fish communities in small streams

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Plerocercoids of the cestode Diphyllobothrium ditremum in brown trout Salmo trutta: substantial increase in infection after establishment of European minnow Phoxinus phoxinus

This study focuses on plerocercoids of the cestode Diphyllobothrium ditremum in brown trout Salmo trutta from the subalpine lake Øvre Heimdalsvatn in south–central Norway. Salmo trutta was the only fish species in this lake until European minnow Phoxinus phoxinus was registered in 1969. The P. phoxinus population increased substantially in the following years. In contrast with the 1969–1972 period, when plerocercoids of D. ditremum were practically absent in S. trutta, there was a high prevalence and intensity of infection in the 2013 S. trutta samples. Because the life cycle of D. ditremum involves two larval stages, in copepods and salmonids and mature worms in piscivorous birds, such as mergansers and loons, a change in feeding ecology of S. trutta or changes in population densities of copepods, fish or birds might have influenced the infection pattern. No relationships between D. ditremum infection and muscle‐tissue δ¹⁵N signature or Hg concentration were found, indicating that infection is not a result of piscivory or cannibalism. Furthermore, consumption of copepods by S. trutta during summer and autumn was low. On the other hand, the number of piscivorous birds has increased, probably due to the presence of P. phoxinus as a new and numerous prey. An increased number of final D. ditremum hosts may have produced a higher output of cestode eggs, resulting in more infected copepods that in turn are consumed by S. trutta. Indirectly, P. phoxinus may therefore have caused the observed increased infection in S. trutta and thereby imposed further negative effects on S. trutta in high mountain areas.     

The role of alien fish (the centrarchid Micropterus salmoides) in lake food webs highlighted by stable isotope analysis

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Mitigation of acidified salmon rivers – effects of liming on young brown trout Salmo trutta

In southern Norway, 22 acidified rivers supporting anadromous salmonids were mitigated with lime to improve water quality and restore fish populations. In 13 of these rivers, effects on Salmo trutta and Salmo salar densities were monitored over 10–12 years, grouped into age 0 and age ≥ 1 year fish. These rivers had a mean annual discharge of between 4·9 and 85·5 m³ s^?1, and six of them were regulated for hydro‐power production. Salmo salar were lost in six of these rivers prior to liming, and highly reduced in the remaining seven rivers. Post‐liming, S. salar became re‐established in all six rivers with lost populations, and recovered in the seven other rivers. Salmo trutta occurred in all 13 study rivers prior to liming. Despite the improved water quality, both age 0 and age ≥ 1 year S. trutta densities decreased as S. salar density increased, with an average reduction of >50% after 10 years of liming. For age 0 year S. trutta this effect was less strong in rivers where S. salar were present prior to liming. In contrast, densities of S. trutta increased in unlimed streams above the anadromous stretches in two of the rivers following improved water quality due to natural recovery. Density increases of both age 0 and age ≥ 1 year S. salar showed a positive effect of river discharge. The results suggest that the decline in S. trutta density after liming is related to interspecific resource competition due to the recovery of S. salar. Thus, improved water quality through liming may not only sustain susceptible species, but can have a negative effect on species that are more tolerant prior to the treatment, such as S. trutta.     

Community structure affects trophic ontogeny in a predatory fish

While most studies have focused on the timing and nature of ontogenetic niche shifts, information is scarce about the effects of community structure on trophic ontogeny of top predators. We investigated how community structure affects ontogenetic niche shifts (i.e., relationships between body length, trophic position, and individual dietary specialization) of a predatory fish, brown trout (Salmo trutta). We used stable isotope and stomach content analyses to test how functional characteristics of lake fish community compositions (competition and prey availability) modulate niche shifts in terms of (i) piscivorous behavior, (ii) trophic position, and (iii) individual dietary specialization. Northern Scandinavian freshwater fish communities were used as a study system, including nine subarctic lakes with contrasting fish community configurations: (i) trout‐only systems, (ii) two‐species systems (brown trout and Arctic charr [Salvelinus alpinus] coexisting), and (iii) three‐species systems (brown trout, Arctic charr, and three‐spined sticklebacks [Gasterosteus aculeatus] coexisting). We expected that the presence of profitable small prey (stickleback) and mixed competitor–prey fish species (charr) supports early piscivory and high individual dietary specialization among trout in multispecies communities, whereas minor ontogenetic shifts were expected in trout‐only systems. From logistic regression models, the presence of a suitable prey fish species (stickleback) emerged as the principal variable determining the size at ontogenetic niche shifts. Generalized additive mixed models indicated that fish community structure shaped ontogenetic niche shifts in trout, with the strongest positive relationships between body length, trophic position, and individual dietary specialization being observed in three‐species communities. Our findings revealed that the presence of a small‐sized prey fish species (stickleback) rather than a mixed competitor–prey fish species (charr) was an important factor affecting the ontogenetic niche‐shift processes of trout. The study demonstrates that community structure may modulate the ontogenetic diet trajectories of and individual niche specialization within a top predator.     

Marine nutrient contributions to freshwater apex predators

Recent investigations into the nutrient cycling of coastal ecosystems have suggested that migratory or anadromous fish could be important vectors of marine nutrients. Anadromous fish have assimilated marine nutrients that would contribute to the nutrient budgets of freshwater systems by excretion, gamete release, or the decay of post-reproductive carcasses. However, the extent to which freshwater predators utilize marine material is not well understood. In systems where anadromous fish temporarily constitute a major portion of the fish community, they may contribute substantially to the diet of piscivorous fish and other predators. Here we show the contribution of anadromous blueback herring, shad, and alewife (Alosa) to diets of large, non-indigenous piscivorous catfish (Ictalurus furcatus) using δ³⁴S and δ¹³C. The spawning anadromous Alosa, captured in tidal freshwater, had enriched δ³⁴S and δ¹³C values compared to resident, native freshwater species. As a result of consuming the anadromous Alosa, the I. furcatus isotope signature shifted towards the marine signal. The isotope analysis revealed that anadromous fish contribute substantially to the diet of most captured I. furcatus. The percentage of anadromous Alosa carbon and sulfur that was incorporated into I. furcatus (≥38 cm total length) ranged from 0 to 84% and 10 to 69%, and had means (±SD) of 42 (±24) and 43 (±16)%, respectively. Although the δ¹⁵N signal of marine-derived biomass is enriched by approximately 3‰ relative to terrestrial or freshwater biomass, it was not as useful as δ³⁴S and δ¹³C for nutrient source owing to trophic fractionation. This study demonstrates that anadromous fish may be a significant source of nutrients to tidal freshwater apex predators. Received: 19 March 1999 / Accepted: 26 October 1999     

The decapod researcher’s guide to the galaxy of sex determination

The objective of this study was to assess the frequency of use of marine resources in recruitment of Southern Hemisphere native riverine fish Galaxias maculatus from rivers across a latitudinal gradient. To do this, we analysed the concentrations of δ³⁴S in vertebral column tissues from fish collected in ten Chilean river systems across latitudes 36°–47°S. The analyses of δ³⁴S signatures in these rivers suggest that the use of marine resources by riverine populations of G. maculatus in large river systems in Chile is variable, with marine resources playing a limited role in more northern large rivers, characterised by warmer temperatures and predictable flow regimes and floodplain inundations. This is in contrast to life histories described for G. maculatus in rivers from New Zealand and Australia, where riverine populations are believed to be characterised by an obligatory recruitment phase in marine environments. Recruitment of G. maculatus in Chilean large rivers appears to depend on their freshwater productivity driven by climate as well as both longitudinal (headwaters lakes-estuary) and lateral (main channel-floodplains) hydrologic connectivities.     

Brown trout and food web interactions in a Minnesota stream

1. We examined indirect, community‐level interactions in a stream that contained non‐native brown trout (Salmo trutta Linnaeus), native brook trout (Salvelinus fontinalis Mitchill) and native slimy sculpin (Cottus cognatus Richardson). Our objectives were to examine benthic invertebrate composition and prey selection of fishes (measured by total invertebrate dry mass, dry mass of individual invertebrate taxa and relative proportion of invertebrate taxa in the benthos and diet) among treatments (no fish, juvenile brook trout alone, juvenile brown trout alone, sculpin with brook trout and sculpin with brown trout). 2. We assigned treatments to 1 m² enclosures/exclosures placed in riffles in Valley Creek, Minnesota, and conducted six experimental trials. We used three designs of fish densities (addition of trout to a constant number of sculpin with unequal numbers of trout and sculpin; addition of trout to a constant number of sculpin with equal numbers of trout and sculpin; and replacement of half the sculpin with an equal number of trout) to investigate the relative strength of interspecific versus intraspecific interactions. 3. Presence of fish (all three species, alone or in combined‐species treatments) was not associated with changes in total dry mass of benthic invertebrates or shifts in relative abundance of benthic invertebrate taxa, regardless of fish density design. 4. Brook trout and sculpin diets did not change when each species was alone compared with treatments of both species together. Likewise, we did not find evidence for shifts in brown trout or sculpin diets when each species was alone or together. 5. We suggest that native brook trout and non‐native brown trout fill similar niches in Valley Creek. We did not find evidence that either species had an effect on stream communities, potentially due to high invertebrate productivity in Valley Creek.     

Interactions between invading benthivorous fish and native whitefish in subarctic lakes

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Trophic interference by Salmo trutta on Aplochiton zebra and Aplochiton taeniatus in southern Patagonian lakes

The length and mass ratio, diet and isotopic composition of Aplochiton zebra and Aplochiton taeniatus inhabiting a Salmo trutta‐invaded and a S. trutta‐free lake in southern Patagonia were compared. Results indicate that S. trutta exercises important trophic interference over A. zebra and A. taeniatus, causing changes in their dietary composition by reducing the consumption of winged Diptera through changes in feeding behaviours that involve jumping out of the water. This effect is significantly higher in A. zebra than in A. taeniatus a species that has a highly specialized diet. The dietary changes of A. zebra and A. taeniatus in sympatry with S. trutta lead to an impoverishment of their isotopic nitrogen signals (δ¹⁵N), suggesting a reduction of their trophic position. In the case of A. zebra, this translates into a significant decrease in its body condition factor. Such interference could lead to a population decline of this species and would explain the current distribution range decline and allopatry with S. trutta in fluvial systems.     

Feeding habits and seasonal trophic guilds structuring fish community in the bay mouth region of a tropical estuarine habitat

Understanding trophic relationships of fish in estuarine ecosystem is an important element for sustainable resource management. This study examined the feeding habits of 29 dominant fish species, characterized the trophic guilds, assessed the impact of season and clarified the role of diets in structuring the fish community in the mouth region of Pattani Bay, Thailand. Samples of 5792 fishes collected monthly by gillnets from March 2019 to February 2020 were used for stomach content analyses. It was found that the number of food types and fullness index differed between fish taxa (P < 0.001). Most fishes were specialist feeders feeding on specific food components and were categorized into five trophic guilds: piscivore, shrimp-fish feeder, polychaete feeder, zooplanktivore and planktivore. Six species were piscivorous, considered as apex predators, that fed almost entirely on fishes. High diet overlaps among some species (>0.6) were recorded. Not much variation in seasonal guilds was observed: four guilds in the dry season, three in the moderate rainy season and four in the rainy season. Some species remained in the same guild the whole year round, but some fishes changed seasonally. Two fish communities from different regions of the bay were segregated based on feeding habits. The inner bay community comprised mainly copepod and plankton feeders, but there were more piscivores in the deeper bay mouth area. Results from this study help us to understand the feeding habits and trophic guilds of dominant fish species at the mouth of this tropical estuarine bay.     

Discrimination between farmed and free-living invasive salmonids in Chilean Patagonia using stable isotope analysis

In Chilean Patagonia relatively pristine aquatic environments are being modified by the introduction of exotic salmonids, initially through their deliberate release for sport fishing since the early twentieth century, and more recently via the accidental escape from fish farms. There is therefore a need to reliably distinguish between naturally reproducing and fugitive salmonids associated with the Chilean salmonid farming industry, the second largest in the world. We tested the ability of stable isotope analysis (SIA) and analysis of scale growth profiles to discriminate between farmed and free-living salmonids sampled around the Island of Chiloé. Juvenile Atlantic salmon (Salmo salar) and rainbow trout (Oncorhynchus mykiss) from aquaculture facilities were significantly more enriched in δ¹⁵N and lipid-corrected δ¹³C than river-caught individuals. Scale growth slopes during the first year in freshwater were significantly higher in farmed than in wild-caught rainbow trout, indicating faster somatic growth under hatchery conditions. Stable isotopes analysis classified 94% of juvenile Atlantic salmon and rainbow trout to their correct farm or free-living groups. Our results, therefore, can help to elucidate the origin and spread of exotic invasive salmonids in Chile, and address one of the biggest threats to native freshwater fishes in Patagonia and other temperate zones of the Southern Hemisphere.     

Differences in growth between offspring of anadromous and freshwater brown trout Salmo trutta

In this study, individual growth of juvenile offspring of anadromous and freshwater resident brown trout Salmo trutta and crosses between the two from the River Imsa, Norway, was estimated. The juveniles were incubated until hatching at two temperatures (±S.D. ), either 4.4 ± 1.5°C or 7.1 ± 0.6°C. Growth rate was estimated for 22 days in August–September when the fish on average were c. 8 g in wet mass, and the estimates were standardized to 1 g fish dry mass. Offspring of anadromous S. trutta grew better at both 15 and 18°C than offspring of freshwater resident S. trutta or offspring of crosses between the two S. trutta types. This difference appears not to result from a maternal effect because anadromous S. trutta grew better than the hybrids with anadromous mothers. Instead, this appears to be an inherited difference between the anadromous and the freshwater resident fish lending support to the hypothesis that anadromous and freshwater resident S. trutta in this river differ in genetic expression. Egg incubation temperature of S. trutta appeared not to influence the later growth as reported earlier from the studies of Atlantic salmon Salmo salar.     

Fish community structure and environmental correlates in the highly altered southern Sacramento-San Joaquin Delta

We sampled 11 sites in the southern Sacramento-San Joaquin Delta from 1992–1999, to characterize fish communities and their associations with environmental variables. Riparian habitats were dominated by rock-reinforced levees, and large water diversion facilities greatly influenced local hydrodynamics and water quality. We captured 33 different taxa, only eight of which were native. None of the native species represented more than 0.5% of the total number of individuals collected. The abundance of native species was consistently low but typically peaked during high outflow periods. Fish communities were predominantly structured along environmental gradients of water temperature and river flow. Native species (tule perch, Hysterocarpus traski, & Sacramento sucker, Catostomus occidentalis) were associated with conditions of high river flow and turbidity, while the majority of the non-native species were associated with either warm water temperature or low river flow conditions. The exceptions were the non-native striped bass, Morone saxatilis, and white catfish, Ameiurus catus, which were positively associated with relatively high river flow. Variation in fish community structure was greater among river locations within years than within river locations among years, thus fish communities at each river location were consistently different each year. Differences in fish communities among river locations were correlated with river flow and turbidity. We predict that the fish communities of this region will remain numerically dominated by non-native species if the environmental conditions we observed persist in the future.     

Response of the fish community to human-induced changes in the Biobío River in Chile

1. The Biobío River basin of south‐central Chile exhibits the greatest species richness of all rivers in Chile, where it is one of the most important rivers for human use. Use for provision of drinking water, irrigation, sewage effluents, hydropower generation and industry has increased dramatically during the last decade. To help understand the effects of human activities on the Biobío River, we document recent changes in the fish community. 2. In this study, current patterns of distribution and abundance of fishes were compared with the expected longitudinal pattern, and to historical data from studies conducted before the rapid development of the last decade. Fish distribution, biomass, abundance and diversity were studied at eight sampling stations in the middle and lower zones of the river in both high and low flow seasons. 3. Contrary to the pattern observed in less impacted river systems, species richness (S), diversity (H′) and abundance [calculated catch per unit effort (CPUE)] all tended to decrease downstream from the uppermost sampling locations. Mean S decreased from 7.9 to 5.4, mean H′ decreased from 0.7 to 0.4, and mean CPUE decreased from 111 to 43 from hyporithral to potamal locations. 4. Comparison with previous records indicates loss or reduction in distribution of native species, and a concurrent expansion in distribution and abundance of tolerant introduced species (e.g. Gambusia holbrooki, and Cyprinus carpio) over the last 10–15 years. These comparisons suggest a large‐scale and long‐term effect of recent human impacts on the river.     

Carbon sources and trophic structure in a macrophyte‐dominated polyculture pond assessed by stable‐isotope analysis

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