Empirical predictions of the trophic consequences of non-native freshwater fishes: a synthesis of approaches and invasion impacts

Predicting the ecological impacts of invasive fishes is crucial for understanding their risks in the environment. Experimental approaches that provide empirical data on invasion impacts provide both theoretical insights and data of high management utility for prioritising high-risk invaders. Here, a synthesis of some experimental approaches that predict invasion impacts of non-native fish is presented, where the focus is on impacts relating to the trophic impacts of the invader on either native trophically analogous fishes or prey populations. Experiments in tank aquaria are advantageous in providing homogenous and controlled conditions that also enable high replication. Competition-based studies can determine invasion impacts via exposure of fish to fixed food rations, although these experiments tend to produce results in intensely competitive conditions that might not occur in the wild. Comparative functional response experiments in tank aquaria have been successfully applied to determining the relative impacts of invaders compared with native species on prey populations, and have revealed high resource efficiency in globally invasive piscivorous fishes. Experiments completed in mesocosms and small ponds have the advantage of providing replicated systems in semi-natural conditions. They have been successful in revealing that rather than competing with native fishes, many invasive fishes show strong patterns of trophic niche partitioning. These patterns are also evident in these fishes when in natural invaded communities. Thus, whilst it is often suggested invasion impacts from non-native fishes result from inter-specific competition, this is being increasingly challenged by a combination of experimental and field-based predictions

Do trophic subsidies affect the outcome of introductions of a non-native freshwater fish?

Understanding how environmental variables and human disturbances influence the outcomes of introductions of non-native freshwater fish is integral to their risk management. This can be complex in freshwater ecosystems that receive subsidies that increase food availability, as these may influence the outcome of introductions through promoting the survival, reproduction and establishment of the introduced propagules through increasing their access to food resources. We determined how natural and/or artificial trophic subsidies affected the reproduction and establishment of the introduced topmouth gudgeon (Pseudorasbora parva) in replicated pond mesocosms. The mesocosms all started with eight mature fish and were run for 100 days during their reproductive season. The subsidies consisted of natural terrestrial prey and/or fishmeal pellets (a common trophic subsidy that can be significant in systems that are used as sport fisheries or for aquaculture). After 100 days, fish in the natural subsidy ponds showed minimal growth and very low reproductive output. Analysis of δC and δN indicated that their progeny, 0+ fish produced in the ponds, exploited the terrestrial prey. By contrast, in ponds where pellets were added, mineral nutrient availability and primary production were significantly increased, and the mature fish fed mainly on the aquatic resources. The increased productivity of the ponds significantly increased fish growth and fitness, resulting in high numbers of 0+ individuals that did feed on the pellets. Thus, subsidies that can increase both primary production and food resources (such as pelletised fishmeal) can significantly influence the ability of colonists to establish a population rapidly. Management efforts to minimise the risk of introductions should thus consider the role of these types of allochthonous subsidies. © 2013 The Authors. Freshwater Biology published by John Wiley & Sons Ltd

Stable isotope discrimination factors of omnivorous fishes: influence of tissue type, temperature, diet composition and formulated feeds

Application of stable isotope data to trophic studies requires understanding of factors influencing the isotopic discrimination factor (Δ) between consumers and their prey resources. This is missing for many omnivorous species, despite their diet and environment potentially impacting Δ. The effects of temperature, diet (including formulated feeds) and tissue type on Δ13C and Δ15N were thus tested experimentally. A temperature experiment exposed three species to identical diets at 18 and 23°C, whereas a diet experiment exposed one species to four diets at 18°C. At 23°C, C:N ratios, Δ13C and Δ15N were generally elevated versus 18°C. After lipid correction, tissue/species-specific differences at 23°C in Δ13C and Δ15N were up to 0.73 and 0.54‰ higher, respectively. Across the four diets, there were also significant differences in Δ13C and Δ15N between a natural diet and diets based on formulated feeds. Δ13C and Δ15N of muscle were 1.51 to 2.76‰ and 3.13 to 5.44‰, respectively. Highest Δ for both isotopes was from a formulated feed based on plant material that resulted in lower dietary protein content and quality. Thus, diet and environment fundamentally affected the isotopic discrimination factors and these factors require consideration within trophic studies based on stable isotopes

A Case Study of the Population Ecology of a Topmouth Gudgeon (Pseudorasbora parva) Population in the UK and the Implication for Native Fish Communities

1. The topmouth gudgeon Pseudorasbora parva is a small Asian cyprinid species that has proved invasive throughout many European countries. Following an initial introduction into the wild in 1996, the species is now proving invasive in the UK, with at least 25 infested waters in England and Wales, of which 10 are known to have direct connection to a major river catchment. 2. To demonstrate the threat of P. parva to fisheries in the UK, a case study is presented on a lake located in the Lake District of England where the species was introduced in 2000. The species rapidly established a breeding population that, by 2003, was the dominant species in size classes <70 mm. In 2004, they were the only species in the lake that produced young-of-the-year. 3. Individual P. parva adopted the reproductive tactics of early maturity, multiple spawning, male dominance and male nest guarding; sexual dimorphism was manifested in larger body size of males. These traits were in contrast to the resident, native species of the lake, including roach Rutilus rutilus and gudgeon Gobio gobio, which adopted traits of later maturity and single spawning. 4. This case study, therefore, revealed relatively rapid establishment of a P. parva population, their subsequent numerical dominance of the fish community, and the impediment of the recruitment of native fish. The implications for UK fisheries are concerning: should P. parva continue to disperse and individuals adopt similar traits as those in this case study, there may be few waters immune from their invasion, numerical dominance and subsequent impacts

Individual variability in stable isotope turnover rates of epidermal mucus according to body size in an omnivorous fish.

Epidermal mucus (‘mucus’) is increasingly applied to fish ecological studies based on stable isotope analysis (SIA) due to its non-invasive collection. However, knowledge on mucus SI turnover rates of individual fish remains limited, including uncertainty over how they are influenced by fish body sizes. Here, a diet switch experiment predicted mucus SI turnover rates (δ13C and δ15N) as a function of time using samples taken over 200 days from 10 individually tagged common carp Cyprinus carpio covering two size groups. Non-linear mixed effects models revealed rapid turnover of both δ13C and δ15N (T50: 2–5 days; T95: 9–22 days); δ15N turnover rates were slower for the larger cohort, while δ13C turnover rates were independent of body size. Within size groups, turnover rates were not expected to vary between individuals. These experimental results suggest that due to these fast turnover rates, epidermal mucus can provide insights into the diets of fish over very short timeframes, although for δ15N the body size of the fish needs consideration

Assessing the efficacy and ecology of biocontrol and biomanipulation for managing invasive pest fish

1. Management of non-native species aims to prevent biological invasions using actions including control and containment of the potential invader. Biocontrol and biomanipulation strategies are used frequently to reduce population sizes of non-native species, and reduce their ecological impacts and dispersal rates. 2. Assessments of the efficacy of biocontrol and biomanipulation actions for managing non-native pest fish, and the ecological mechanisms involved, were studied here using lentic populations of the invasive fish Pseudorasbora parva. Biocontrol was through release of the indigenous piscivorous fish Perca fluviatilis and biomanipulation through intensive fish removals. 3. A combined biocontrol and removal programme was completed in an invaded pond over 2 reproductive seasons. Almost 10000 P. parva were removed, with cumulative removal numbers significantly related to their decreased abundance (>60 to <0.1 m-2). Ten adult P. fluviatilis were also released initially and reproduced each season. Analyses revealed P. parva contribution to their diet was high initially, but decreased as P. parva abundance reduced. Individual contributions of the management actions to declined P. parva abundance were difficult to isolate. 4. The individual effects of biocontrol and removals on P. parva populations were then tested using a field trial in replicated pond mesocosms over 3 reproductive seasons. Replicates started with 1500 P. parva. The control (no interventions) revealed no significant temporal changes in P. parva abundances. In the removal treatment, where over 17000 P. parva were removed per replicate over the trial, abundance declined initially, but increased significantly after each reproductive season as remaining fish compensated through increased reproductive output. In the biocontrol, abundance declined and remained low; analyses revealed P. parva were an important dietary component of larger P. fluviatilis, with predation suppressing compensatory responses. 5. Synthesis and applications: Biocontrol and removals can significantly reduce abundances of lentic populations of small invasive fishes. Removals provide short-term population suppression, but high effort is needed to overcome compensatory responses. Biocontrol can provide longer-term suppression but could invoke unintended ecological consequences via ‘stocking-up’ food webs. Application of these results to decision-making frameworks should enable managers to make more objective decisions on risk-commensurate methodologies for controlling small invasive fishes

Spawning strategies in cypriniform fishes in a lowland river invaded by non-indigenous European barbel Barbus barbus

© 2020, The Author(s). Spawning strategies of lowland river fishes include single spawning, where reproduction generally occurs in early spring to provide 0+ fish with an extended growth season through the summer, but with a high risk of stochastic mortality events occurring, such as early summer floods. This risk can be reduced by multiple or protracted spawning strategies, where 0+ fish are produced over an extended period, often into mid-summer, but with the trade-off being a shorter growth season. The spawning strategies of cypriniform fish were explored in the River Teme, a spate river in Western England, which has non-indigenous European barbel Barbus barbus present. Sampling 0+ fish in spring and summer and across three spawning periods, B. barbus, chub Squalius cephalus and minnow Phoxinus phoxinus always revealed multiple spawning events, with 0+ fish of < 20 mm present in samples collected from June to August. Fish below 20 mm in August remained relatively small by the end of their growth season (October). For dace Leuciscus leuciscus, only single spawning events were evident, but with 0+ dace always being relatively large. Therefore, multiple spawning appears to be a common strategy that provides resilience in 0+ fish against stochastic mortality events in lowland rivers

Sea Lamprey Petromyzon marinus Biology and Management Across Their Native and Invasive Ranges: Promoting Conservation by Knowledge Transfer

The anadromous sea lamprey Petromyzon marinus native range extends across the Northern Atlantic and includes much of Europe. Their complex lifecycle, involving freshwater spawning, juveniles (ammocoetes) that remain in freshwater for up to eight years, and adults migrating to sea before returning to reproduce, means native populations in Europe are threatened by multiple stressors, especially migration blockages and habitat loss. This has resulted in population declines across their European range, despite their ecological, evolutionary, and economic significance. Information on their population demography and long-term patterns are also scarce, with focus primarily on their ammocoete freshwater phase. This is inhibiting the development of biological reference points for utilization in population monitoring programs. In the Great Lakes of North America, however, P. marinus is invasive and the high damage caused to commercial fisheries resulted in their populations being controlled through a long-term, multi-method and integrated research and management approach over the last 40 years, with the development and application of a range of novel methods. Successful knowledge transfer to Europe could therefore facilitate the monitoring of threatened populations and develop new conservation actions, including modifying migration blockages to facilitate passage, implementing adult trapping programs, and applying pheromone treatments to manipulate adult movements and behaviors. This reveals the potential utility of using invasive fish populations to inform conservation practices in native ranges, and how pheromone research could further enhance fish conservation and monitoring

Spatial variability in the growth of invasive European barbel Barbus barbus in the River Severn basin, revealed using anglers as citizen scientists

Life history trait analyses of non-native fishes help identify how novel populations respond to different habitat typologies. Here, using electric fishing and anglers as citizen scientists, scales were collected from the invasive barbel Barbus barbus population from four reaches of the River Severn and Teme, western England. Angler samples were biased towards larger fish, with the smallest fish captured being 410 mm, whereas electric fishing sampled fish down to 60 mm. Scale ageing revealed fish present to over 20 years old in both rivers. Juvenile growth rates were similar across all reaches. Lengths at the last annulus and Linfinity of the von Bertalanffy growth model revealed, however, that fish grew to significantly larger body sizes in a relatively deep and highly impounded reach of the River Severn. Anglers thus supplemented the scale collection and although samples remained limited in number, they provided considerable insights into the spatial demographics of this invasive B. barbus population