were that the invaders would have significantly higher consumption rates for 34 chironomids than the native fishes, but not for G. pulex. Mean consumption rates for 35 chironomids were significantly lower for both invaders than B. barbus and S. 36 cephalus, but were similar to T. tinca. Barbus barbus had a significantly lower 37 consumption rate of G. pulex than both invaders, but there were no significant 38 differences between S. cephalus, T. tinca and the invaders. All FRs were type II, with 39 functional response curves for the invaders preying upon chironomids never being 40 significantly higher than the native fishes, contrary to predictions. For G. pulex, some 41 significant differences were apparent between the invaders and native fishes, but 42 again were contrary to predictions. These results indicated that when predation 43 impacts of invasive fishes could also be a function of their population density and 44 body sizes, these parameters should be incorporated into FR models to improve 45 impact predictions. 46
The crossing of freshwater ecosystem boundaries by marine‐derived nutrients (MDN) is usually associated with migratory salmonid fishes returning to natal rivers. An alternative source of MDN in fresh waters is the widespread use of pelletised marine fishmeal (“pellets”) by freshwater anglers as they target large‐bodied cyprinid fishes, such as European barbel Barbus barbus. Here, the trophic consequences of MDN from pellets for riverine cyprinid fishes were tested. Approaches used stable isotope analyses in controlled and wild scenarios, using B. barbus and chub Squalius cephalus as model species. The isotopic niche, measured as standard ellipse area, was used to assess trophic niche size, and mixing models predicted the extent to which MDN contributed to fish diet. In experimental mesocosms, B. barbus fed low volumes of pellets (c. 3 per fish) for 130 days had isotopic niche sizes that were up to four times larger than a control and “medium” (6 per fish) and “high” pellet (12 per fish) treatments. Somatic growth rates were significantly higher in the “medium” and “high” treatments. In pond enclosure experiments, when juvenile B. barbus and S. cephalus were fed pellets daily for 100 days, there was a substantial and significant shift in the position of their isotopic niche compared to controls with no pellets fed. However, for each species, there were no significant differences in their somatic growth rates in the presence/absence of pellets. In a lowland river, high proportions of MDN contributed to the diet of B. barbus and S. cephalus captured by angling, but with substantial individual variability in those captured by electric fishing. Across all B. barbus >400 mm, MDN dietary contributions ranged between 9% and 71%. This suggested some individual diet specialisations within their population that was associated with feeding on this angler subsidy and that also resulted in a significant increase in the size of their population isotopic niche. These results suggested that when pellets containing MDN are used in freshwater angling, they are consumed and assimilated by cyprinid fishes, influencing individual and population trophic positions, and isotopic niche sizes and dietary specialisations. The results also suggested that the extent to which individuals specialise in feeding on pellets potentially influences their vulnerability to capture by anglers.
Ecological theory on the trophic impacts of invasive fauna on native competitors is equivocal. Whilst increased interspecific competition can result in coexisting species having constricted and diverged trophic niches, the competing species might instead increase their niche sizes to maintain energy intakes. Empirical experiments can test invasion theory on competitive interactions and niche sizes across different spatial scales and complexity. The consequences of increased interspecific competition from a model alien fish Leuciscus idus were tested on two taxonomically and trophically similar native fishes, Squalius cephalus and Barbus barbus. Competitive interactions were tested in tank aquaria using comparative functional responses (CFRs) and cohabitation trials. The consequences of these competitive interactions for the trophic niche sizes and positions of the fishes were tested in pond mesocosms. Comparative functional responses revealed that compared to B. barbus, L. idus had significantly higher attack and consumption rates; cohabitation trials revealed B. barbus growth rates were depressed in sympatry with L. idus. For L. idus and S. cephalus, differences in their functional response parameters and growth rates were not significant. Pond mesocosms used stable isotope metrics to quantify shifts in the trophic niche sizes of the fishes between allopatry and sympatry using a substitutive experimental design. Isotopic niches were smaller and more divergent in sympatric paired species than predicted by their allopatric treatments, suggesting trophic impacts from interspecific competition. However, an all‐species sympatric treatment revealed similar niche sizes with allopatry. This maintenance of niche sizes in the presence of all species potentially resulted from the buffering of direct competitive effects of the species pairs by indirect effects. Experimental predictions from tank aquaria assisted the interpretation of the constricted and diverged trophic niches detected in the paired‐species sympatric treatments of the pond mesocosms. However, the all‐species sympatric treatment of this experiment revealed greater complexity in the outcomes of the competitive interactions within and between the species. These results have important implications for understanding how alien species integrate into food webs and influence the trophic relationships between native species.
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