Selective predators in complex communities – mechanisms and consequences of benthic fish predation in small temperate streams

The prey consumption by benthivorous fish predators can have profound top-down effects in stream food webs. To analyse this effects in small temperate stream ecosystems, a long-term field experiment was conducted in two streams in South-eastern Germany, Gauernitzbach and Tännichtgrundbach, from 2004 to 2011. The densities of two small-bodied benthivorous fish species, gudgeon (Gobio gobio) and stone loach (Barbatula barbatula), were manipulated following a Before-After-Control-Impact design. The top predator regime affected the benthic community composition of the streams mainly in pools, whereas the total benthic invertebrate biomass was not affected in any mesohabitat. The present work describes a causal analysis of the observed food web effects using additional field analyses and laboratory experiments, with a special focus on the habitat use and foraging behaviour of the fish as top predators. The probably most important of the analysed mechanisms was mesohabitat-specific predation by the fish. Three 24-h field video surveys combined with benthic invertebrate sampling showed that constraints in habitat use, especially for gudgeon, induced a differential predator-prey habitat overlap which resulted in a higher predation risk for the invertebrate prey in pools than in riffles. Another important mechanism was selective predation of both fish species. Their prey selectivity was largely explained by a small number of prey variables being connected to the partly non-visual foraging mode of these benthic predators. Besides the traits body size and feeding type, long-term mean abundance played a central role, small and highly abundant invertebrates, grazers and sediment feeders being preferred by gudgeon and stone loach. The preference for small and abundant prey taxa (chironomids) exceeded purely opportunistic feeding and probably facilitated resource partitioning between the two fish species having very similar diets. In addition to active selectivity, different predator avoidance strategies of the invertebrates analysed in laboratory experiments explained the passive selectivity of the fish predators for certain prey taxa in the streams. This could be shown for two abundant taxa being consumed by the fish predators in very different quantities, Gammarus pulex and Hydropsyche instabilis. These three mechanisms, although probably interacting with several other factors, could explain a large part of the effects the top-down food web manipulation had on the benthic community, especially the observed high degree of mesohabitat and species specificity. Confirming this, quantitative characteristics of predation food webs, for instance the importance of intraguild predation, differed markedly between pool and riffle mesohabitats. From the results of this study it can be concluded that the benthivorous fish affected benthic community structure mainly by mesohabitat-specific and selective predation. A manipulation of this (native) top predator type therefore will probably have such rather subtle but not catastrophic consequences in ecosystems with a high biotic diversity and a rich natural habitat structure and connectivity

Extended larval development compensates for sublethal effects of fish predation in a mayfly population (Rhithrogena semicolorata, Ephemeroptera)

AbstractMany predation experiments in streams are carried out in enclosures. Hence, the relevance of their results to predict population dynamics is often unclear due to the relatively small spatial and temporal scale of the experiments. To enhance the transferability of experimental results on the ecosystem scale the impact of fish predators on a prey population was observed in a reach scale approach over 2 years in a natural stream. A 400-m reach inhabited by the small benthivorous fishes gudgeon (Gobio gobio) and stone loach (Barbatula barbatula) was compared with a fishless reference reach.It was shown that fish predation may affect the population of the grazing mayfly Rhithrogena semicolorata on the ecosystem scale. Although the larvae grew slower in the fish reach than in the fishless reach, the adults reached the same size and fecundity because they emerged 2–3 weeks later. By this compensation, the prey species avoided a reduction of their individual fecundity. On the other hand, the extended exposure to the fish predators resulted in an enhanced mortality and a reduced density of adult mayflies. Thus, there was obviously a trade-off between maximising fecundity and minimising mortality from fish predation.The observed differences were almost certainly caused be fish predation and not by natural differences of the reaches. This was concluded from results gained after eliminating all benthivorous fish from the former fish reach.With the help of scenario analyses based on our empirical data and simple model assumptions we could demonstrate that compensating the potential loss in fecundity by extending development time led to higher average fitness of the prey population than emerging at an earlier fixed time to avoid additional predation losses. Therefore, we concluded that this strategy was adaptive in the presence of benthivorous fish

A global agenda for advancing freshwater biodiversity research

Global freshwater biodiversity is declining dramatically, and meeting the challenges of this crisis requires bold goals and the mobilisation of substantial resources. While the reasons are varied, investments in both research and conservation of freshwater biodiversity lag far behind those in the terrestrial and marine realms. Inspired by a global consultation, we identify 15 pressing priority needs, grouped into five research areas, in an effort to support informed stewardship of freshwater biodiversity. The proposed agenda aims to advance freshwater biodiversity research globally as a critical step in improving coordinated actions towards its sustainable management and conservation

A global agenda for advancing freshwater biodiversity research

Global freshwater biodiversity is declining dramatically, and meeting the challenges of this crisis requires bold goals and the mobilisation of substantial resources. While the reasons are varied, investments in both research and conservation of freshwater biodiversity lag far behind those in the terrestrial and marine realms. Inspired by a global consultation, we identify 15 pressing priority needs, grouped into five research areas, in an effort to support informed stewardship of freshwater biodiversity. The proposed agenda aims to advance freshwater biodiversity research globally as a critical step in improving coordinated actions towards its sustainable management and conservation.Peer reviewe

Selective predators in complex communities – mechanisms and consequences of benthic fish predation in small temperate streams

The prey consumption by benthivorous fish predators can have profound top-down effects in stream food webs. To analyse this effects in small temperate stream ecosystems, a long-term field experiment was conducted in two streams in South-eastern Germany, Gauernitzbach and Tännichtgrundbach, from 2004 to 2011. The densities of two small-bodied benthivorous fish species, gudgeon (Gobio gobio) and stone loach (Barbatula barbatula), were manipulated following a Before-After-Control-Impact design. The top predator regime affected the benthic community composition of the streams mainly in pools, whereas the total benthic invertebrate biomass was not affected in any mesohabitat. The present work describes a causal analysis of the observed food web effects using additional field analyses and laboratory experiments, with a special focus on the habitat use and foraging behaviour of the fish as top predators. The probably most important of the analysed mechanisms was mesohabitat-specific predation by the fish. Three 24-h field video surveys combined with benthic invertebrate sampling showed that constraints in habitat use, especially for gudgeon, induced a differential predator-prey habitat overlap which resulted in a higher predation risk for the invertebrate prey in pools than in riffles. Another important mechanism was selective predation of both fish species. Their prey selectivity was largely explained by a small number of prey variables being connected to the partly non-visual foraging mode of these benthic predators. Besides the traits body size and feeding type, long-term mean abundance played a central role, small and highly abundant invertebrates, grazers and sediment feeders being preferred by gudgeon and stone loach. The preference for small and abundant prey taxa (chironomids) exceeded purely opportunistic feeding and probably facilitated resource partitioning between the two fish species having very similar diets. In addition to active selectivity, different predator avoidance strategies of the invertebrates analysed in laboratory experiments explained the passive selectivity of the fish predators for certain prey taxa in the streams. This could be shown for two abundant taxa being consumed by the fish predators in very different quantities, Gammarus pulex and Hydropsyche instabilis. These three mechanisms, although probably interacting with several other factors, could explain a large part of the effects the top-down food web manipulation had on the benthic community, especially the observed high degree of mesohabitat and species specificity. Confirming this, quantitative characteristics of predation food webs, for instance the importance of intraguild predation, differed markedly between pool and riffle mesohabitats. From the results of this study it can be concluded that the benthivorous fish affected benthic community structure mainly by mesohabitat-specific and selective predation. A manipulation of this (native) top predator type therefore will probably have such rather subtle but not catastrophic consequences in ecosystems with a high biotic diversity and a rich natural habitat structure and connectivity

They do not fear the unknown: Ancylus fluviatilis (Mollusca, Planorbidae) shows no predator avoidance behavior towards a novel invasive predator

Biological invasion is a strong threat to native biodiversity, with limnic systems being especially vulnerable due to historical separation and resulting prey naivety. The prey naivety hypothesis states that native species may not be able to recognize novel predators due to a lack of common evolutionary background and, therefore, become easy targets. In a laboratory experiment, we added cues of native European bullhead (Cottus gobio Linnaeus, 1758) and invasive round goby [Neogobius melanostomus (Pallas, 1814)] to Ancylus fluviatilis Müller, 1774 originating from two different populations within the same river (one naive, one experienced towards round goby) and compared their predator avoidance behavior. Individuals from both populations recognized cues from the known predator C. gobio and reduced their locomotive activity. To round goby cues, however, naive individuals did not respond, thereby supporting the prey naivety hypothesis. Experienced individuals, in contrast, reduced their activity, suggesting a learning effect due to the co-occurrence of invasive predator and prey. At fast moving invasion fronts of highly invasive species like N. melanostomus, prey naivety can, hence, enhance their negative impact on ecosystems. Behavioral adaptation of native species resulting in predator avoidance reactions could, therefore, play an important role in ecosystem resilience and temporal invasion dynamics

Round goby

Aquatic biodiversity is threatened by spread of invasive alien species. Round goby Neogobius melanostomus is an invasive fish in large European rivers as well as in coastal waters near their mouths and marbled crayfish Procambarus virginalis is a highly invasive crustacean. Both are small, bottom-dwelling species occupying similar habitat and shelters and utilizing similar food sources. We hypothesized that goby presents a threat to both native and non-native astacofauna in invaded ecosystems. We tested this through laboratory experiments designed to determine aggressiveness and competitiveness of goby against marbled crayfish as a model for other North American cambarid crayfish, assessing goby prey size selection and competition with marbled crayfish for space and shelter. Gobies showed high aggressiveness and dominance over the crayfish. Goby predation on juvenile crayfish was limited by mouth gape size. In goby/crayfish pairs of similar weight, gobies were more aggressive, although each affected the behavior of the other

Diet composition and trophic niche differentiation of Neogobius melanostomus along an invasion gradient in a large lowland river

The Ponto-Caspian goby Neogobius melanostomus (Pallas 1814) is a widespread non-native fish species in European rivers. It occurs in high abundance and can potentially alter the food webs of invaded water bodies profoundly. However, the invasion process usually goes unnoticed, and changes in food webs during the early stages of invasions are rarely described. Here, we studied populations of N. melanostomus along an invasion gradient in the Elbe River (Germany) and tested the effects of time since invasion on the diets and the associated trophic dynamics in two dominant habitats, i.e., rip rap and sand. Results showed that the abundance and biomass of N. melanostomus was lowest at the most recently invaded sites. The trophic niche space quantified by stable isotopes (delta (13) C and delta (15) N) showed that populations at earlier invaded sites exhibited a wider trophic niche than those at the most recently invaded sites. Diptera and Crustacea were the most abundant taxa in N. melanostomus guts at both habitat structures. At rip rap habitats, N. melanostomus showed a significant increasing preference for Crustacea and a decreasing one for Gastropoda along the invasion gradient. At sand habitats no significant relationships between the invasion gradient and electivity index for N. melanostomus were detected. We demonstrated that the invasion stage and habitat in which N. melanostomus occurs affect the feeding preference of this invasive fish and that these factors are therefore important for tracing changes in the macroinvertebrate communities of affected river systems