Non-native fishes and native species diversity in freshwater fish assemblages across the United States

The proliferation of non-native species in North American freshwater ecosystems is considered a primary threat to the integrity of native community structure. However, a general understanding of consistent and predictable impacts of non-native species on native freshwater diversity is limited, in part, because of a lack of broad-scale studies including data from numerous localities across multiple drainages. This study uses data from 751 localities collected during the United States Geological Survey (USGS) National Water Quality Assessment (NAWQA) program to examine the influence of non-native fish species on native freshwater fish assemblages across the United States. In general, no significant differences in native fish richness and diversity measures were detected between sites with only native species and sites containing non-native species. However, at sites with non-native species, the number of non-native species present was negatively correlated with native species richness and Shannon diversity and positively correlated with native evenness. Non-native piscivores were negatively correlated with native species richness and Shannon diversity and positively correlated with native evenness. Native piscivores were positively correlated with native richness and diversity and negatively correlated with native evenness at sites with only native species. Our results suggest that from a superficial perspective, native species richness and diversity are not different among sites with and without non-native species. However, when patterns of native species richness and diversity are examined at sites containing non-native species, correlations between non-native and native species richness and diversity imply the expected negative effect of invasive taxa. Additionally, non-native piscivores appear to have a significant negative effect on native taxa and possibly represent a novel selective force on naive native prey.     

Landscape‐scale determinants of non‐native fish communities

Aim Predicting and preventing invasions depends on knowledge of the factors that make ecosystems susceptible to invasion. Current studies generally rely on non‐native species richness (NNSR) as the sole measure of ecosystem invasibility; however, species identity is a critical consideration, given that different ecosystems may have environmental characteristics suitable to different species. Our aim was to examine whether non‐native freshwater fish community composition was related to ecosystem characteristics at the landscape scale. Location United States. Methods We described spatial patterns in non‐native freshwater fish communities among watersheds in the Mid‐Atlantic region of the United States based on records of establishment in the U.S. Geological Survey’s Nonindigenous Aquatic Species Database. We described general relationships between non‐native species and ecosystem characteristics using canonical correspondence analysis. We clustered watersheds by non‐native fish community and described differences among clusters using indicator species analysis. We then assessed whether non‐native communities could be predicted from ecosystem characteristics using random forest analysis and predicted non‐native communities for uninvaded watersheds. We estimated which ecosystem characteristics were most important for predicting non‐native communities using conditional inference trees. Results We identified four non‐native fish communities, each with distinct indicator species. Non‐native communities were predicted based on ecosystem characteristics with an accuracy of 80.6%, with temperature as the most important variable. Relatively uninvaded watersheds were predicted to be invasible by the most diverse non‐native community. Main conclusions Non‐native species identity is an important consideration when assessing ecosystem invasibility. NNSR alone is an insufficient measure of invasibility because ecosystems with equal NNSR may not be equally invasible by the same species. Our findings can help improve predictions of future invasions and focus management and policy decisions on particular species in highly invasible ecosystems.     

Current knowledge on non‐native freshwater fish introductions

This review provides a contemporary account of knowledge on aspects of introductions of non‐native fish species and includes issues associated with introduction pathways, ecological and economic impacts, risk assessments, management options and impact of climate change. It offers guidance to reconcile the increasing demands of certain stakeholders to diversify their activities using non‐native fishes with the long‐term sustainability of native aquatic biodiversity. The rate at which non‐native freshwater fishes have been introduced worldwide has doubled in the space of 30 years, with the principal motives being aquaculture (39%) and improvement of wild stocks (17%). Economic activity is the principal driver of human‐mediated non‐native fish introductions, including the globalization of fish culture, whereby the production of the African cichlid tilapia is seven times higher in Asia than in most areas of Africa, and Chile is responsible for c. 30% of the world's farmed salmon, all based on introduced species. Consequently, these economic benefits need balancing against the detrimental environmental, social and economic effects of introduced non‐native fishes. There are several major ecological effects associated with non‐native fish introductions, including predation, habitat degradation, increased competition for resources, hybridization and disease transmission. Consideration of these aspects in isolation, however, is rarely sufficient to adequately characterize the overall ecological effect of an introduced species. Regarding the management of introduced non‐native fish, pre‐introduction screening tools, such as the fish invasiveness scoring kit (FISK), can be used to ensure that species are not introduced, which may develop invasive populations. Following the introduction of non‐native fish that do develop invasive populations, management responses are typified by either a remediation or a mitigation response, although these are often difficult and expensive to implement, and may have limited effectiveness.     

Multi-scale phylogenetic structure in coastal dune plant communities across the globe

Aims Studies integrating phylogenetic history and large-scale community assembly are few, and many questions remain unanswered. Here, we use a global coastal dune plant data set to uncover the important factors in community assembly across scales from the local filtering processes to the global long-term diversification and dispersal dynamics. Coastal dune plant communities occur worldwide under a wide range of climatic and geologic conditions as well as in all biogeographic regions. However, global patterns in the phylogenetic composition of coastal dune plant communities have not previously been studied.Methods The data set comprised vegetation data from 18463 plots in New Zealand, South Africa, South America, North America and Europe. The phylogenetic tree comprised 2241 plant species from 149 families. We calculated phylogenetic clustering (Net Relatedness Index, NRI, and Nearest Taxon Index, NTI) of regional dune floras to estimate the amount of in situ diversification relative to the global dune species pool and evaluated the relative importance of land and climate barriers for these diversification patterns by geographic analyses of phylogenetic similarity. We then tested whether dune plant communities exhibit similar patterns of phylogenetic structure within regions. Finally, we calculated NRI for local communities relative to the regional species pool and tested for an association with functional traits (plant height and seed mass) thought to vary along sea–inland gradients.Important findings Regional species pools were phylogenetically clustered relative to the global pool, indicating regional diversification. NTI showed stronger clustering than NRI pointing to the importance of especially recent diversifications within regions. The species pools grouped phylogenetically into two clusters on either side of the tropics suggesting greater dispersal rates within hemispheres than between hemispheres. Local NRI plot values confirmed that most communities were also phylogenetically clustered within regions. NRI values decreased with increasing plant height and seed mass, indicating greater phylogenetic clustering in communities with short maximum height and good dispersers prone to wind and tidal disturbance as well as salt spray, consistent with environmental filtering along sea–inland gradients. Height and seed mass both showed significant phylogenetic signal, and NRI tended to correlate negatively with both at the plot level. Low NRI plots tended to represent coastal scrub and forest, whereas high NRI plots tended to represent herb-dominated vegetation. We conclude that regional diversification processes play a role in dune plant community assembly, with convergence in local phylogenetic community structure and local variation in community structure probably reflecting consistent coastal-inland gradients. Our study contributes to a better understanding of the globally distributed dynamic coastal ecosystems and the structuring factors working on dune plant communities across spatial scales and regions.     

The role of environmental and spatial processes in structuring native and non‐native fish communities across thousands of lakes

Quantifying the role of spatial patterns is an important goal in ecology to further understand patterns of community composition. We quantified the relative role of environmental conditions and regional spatial patterns that could be produced by environmental filtering and dispersal limitation on fish community composition for thousands of lakes. A database was assembled on fish community composition, lake morphology, water quality, climatic conditions, and hydrological connectivity for 9885 lakes in Ontario, Canada. We utilized a variation partitioning approach in conjunction with Moran's Eigenvector Maps (MEM) and Asymmetric Eigenvector Maps (AEM) to model spatial patterns that could be produced by human‐mediated and natural modes of dispersal. Across 9885 lakes and 100 fish species, environmental factors and spatial structure explained approximately 19% of the variation in fish community composition. Examining the proportional role of spatial structure and environmental conditions revealed that as much as 90% of the explained variation in native species assemblage composition is governed by environmental conditions. Conversely on average, 67% of the explained variation in non‐native assemblage composition can be related to human‐mediated dispersal. This study highlights the importance of including spatial structure and environmental conditions when explaining patterns of community composition to better discriminate between the ecological processes that underlie biogeographical patterns of communities composed of native and non‐native fish species.     

Modelling temperature-driven changes in species associations across freshwater communities

Due to global climate change–induced shifts in species distributions, estimating changes in community composition through the use of Species Distribution Models has become a key management tool. Being able to determine how species associations change along environmental gradients is likely to be pivotal in exploring the magnitude of future changes in species’ distributions. This is particularly important in connectivity-limited ecosystems, such as freshwater ecosystems, where increased human translocation is creating species associations over previously unseen environmental gradients. Here, we use a large-scale presence–absence dataset of freshwater fish from lakes across the Fennoscandian region in a Joint Species Distribution Model, to measure the effect of temperature on species associations. We identified a trend of negative associations between species tolerant of cold waters and those tolerant of warmer waters, as well as positive associations between several more warm-tolerant species, with these associations often shifting depending on local temperatures. Our results confirm that freshwater ecosystems can expect to see a large-scale shift towards communities dominated by more warm-tolerant species. While there remains much work to be done to predict exactly where and when local extinctions may take place, the model implemented provides a starting-point for the exploration of climate-driven community trends. This approach is especially informative in regards to determining which species associations are most central in shaping future community composition, and which areas are most vulnerable to local extinctions.     

Non‐native species have multiple abundance–impact curves

The abundance–impact curve is helpful for understanding and managing the impacts of non‐native species. Abundance–impact curves can have a wide range of shapes (e.g., linear, threshold, sigmoid), each with its own implications for scientific understanding and management. Sometimes, the abundance–impact curve has been viewed as a property of the species, with a single curve for a species. I argue that the abundance–impact curve is determined jointly by a non‐native species and the ecosystem it invades, so that a species may have multiple abundance–impact curves. Models of the impacts of the invasive mussel Dreissena show how a single species can have multiple, noninterchangeable abundance–impact curves. To the extent that ecosystem characteristics determine the abundance–impact curve, abundance–impact curves based on horizontal designs (space‐for‐time substitution) may be misleading and should be used with great caution, it at all. It is important for scientists and managers to correctly specify the abundance–impact curve when considering the impacts of non‐native species. Diverting attention from the invading species to the invaded ecosystem, and especially to the interaction between species and ecosystem, could improve our understanding of how non‐native species affect ecosystems and reduce uncertainty around the effects of management of populations of non‐native species.     

Non‐random correlation of species dynamics in tropical tree communities

The importance of environmental stochasticity for tropical tree dynamics has been recently stressed by several studies. This has spurred the development of a ‘time‐averaged neutral model’ of community dynamics by Kalyuzhny and colleagues that extends the neutral model by incorporating environmental stochasticity. We here show that this framework can be used to assess the presence of non‐random correlations between species dynamics. Indeed, the time‐averaged neutral model makes the simplifying assumption that species responses to environmental variation are uncorrelated. We therefore propose to use this model as a null hypothesis against which observed community dynamics can be compared. This study makes five contributions. First, we describe a novel time‐averaged neutral model of community dynamics that is close to, but more flexible than the one previously proposed by Kalyuzhny and colleagues. Second, we develop an inference method based on approximate Bayesian computation (ABC) and demonstrate the identifiability of the model parameters from community time series data. Third, we develop a test of the significance of environmental stochasticity, and a method to quantify its contribution to population variance. Fourth, we develop a test of non‐random correlation between species dynamics. Fifth, we apply these developments to three datasets of tropical tree dynamics. We evidence both a strong contribution of environmental stochasticity to population variance in the three datasets, and a non‐random correlation of species dynamics in one of them. We finally discuss the implications of these results for the modelling of tropical tree community dynamics.     

Non‐native plant species benefit from disturbance: a meta‐analysis

Disturbances, such as fire and grazing, are often claimed to facilitate plant species richness and plant invasions in particular, although empirical evidence is contradictory. We conducted a meta‐analysis to synthesize the literature on how non‐native plant species are affected by disturbances. We explored whether the observed impact of disturbance on non‐native plant communities is related to its type and frequency, to habitat type, study approach (observational or experimental), and to the temporal and spatial scales of the study. To put the results in a broader context, we also conducted a set of parallel analyses on a data set involving native plant species. The diversity and abundance of non‐native plant species were significantly higher at disturbed sites than at undisturbed sites, while the diversity and abundance of native plant species did not differ between the two types of sites. The effect of disturbance on non‐native plant species depended on the measure used to evaluate the impact (species diversity or abundance) and on disturbance type, with grazing and anthropogenic disturbances leading to higher diversity and abundance of non‐native plant species than other disturbance types examined. The impact of disturbance on non‐natives was also associated with study approach, habitat type and temporal scale, but these factors covaried with disturbance type, complicating the interpretation of the results. Overall, our results indicate that disturbance has a positive impact particularly on non‐native plant species (at least when they are already present in the community), and that the strength of this impact depends primarily on the disturbance type. Synthesis Empirical evidence of the effect of disturbances on plant species richness is contradictory. Here we use a meta‐analysis to synthesize the published literature on how different types of disturbances influence the diversity and abundance of plant species, focusing in particular on non‐native plants. Our study supports the hypothesis that disturbances generally facilitate the diversity and abundance of non‐native plant species, although the strength of this facilitation depends primarily on the disturbance type.     

Variable survival across low pH gradients in freshwater fish species

A series of 14 day experiments was conducted on five common New Zealand fish species (redfin bully Gobiomorphus huttoni, inanga Galaxias maculatus, brown trout Salmo trutta, longfin eel Anguilla dieffenbachii and koaro Galaxias brevipinnis) to assess the effect of pH on survival and changes in body mass. No species survived in water of pH <4 although there was 100% survival of all adults at pH 4·5, G. maculatus larvae were also tested and had high mortality at this pH. Results suggest that adults are tolerant of low‐pH waters; however, successful remediation of anthropogenically acidified streams will require an understanding of the susceptibility to low pH on different life cycle stages.     

Non‐parallel divergence across freshwater and marine three‐spined stickleback Gasterosteus aculeatus populations

This work investigated whether multiple freshwater populations of three‐spined stickleback Gasterosteus aculeatus in different freshwater catchments in the Jutland Peninsula, Denmark, derived from the same marine populations show repeated adaptive responses. A total of 327 G. aculeatus collected at 13 sampling locations were screened for genetic variation using a combination of 70 genes putatively under selection and 26 neutral genes along with a marker linked to the ectodysplasin gene (eda), which is strongly correlated with plate armour morphs in the species. A highly significant genetic differentiation was found that was higher among different freshwater samples than between marine–freshwater samples. Tests for selection between marine and freshwater populations showed a very low degree of parallelism and no single nucleotide polymorphism was detected as outlier in all freshwater–marine pairwise comparisons, including the eda. This suggests that G. aculeatus is not necessarily the prime example of parallel local adaptation suggested in much of the literature and that important exceptions exist (i.e. the Jutland Peninsula). While marine populations in the results described here showed a high phenotype–genotype correlation at eda, a low association was found for most of the freshwater populations. The most extreme case was found in the freshwater Lake Hald where all low‐plated phenotypes were either homozygotes for the allele supposed to be associated with completely plated morphs or heterozygotes, but none were homozygotes for the putative low‐plated allele. Re‐examination of data from seven G. aculeatus studies agrees in showing a high but partial association between phenotype–genotype at eda in G. aculeatus freshwater populations and that mismatches occur everywhere in the European regions studied (higher in some areas, i.e. Denmark). This is independent of the eda marker used.     

Soil pathogen communities associated with native and non‐native Phragmites australis populations in freshwater wetlands

Soil pathogens are believed to be major contributors to negative plant–soil feedbacks that regulate plant community dynamics and plant invasions. While the theoretical basis for pathogen regulation of plant communities is well established within the plant–soil feedback framework, direct experimental evidence for pathogen community responses to plants has been limited, often relying largely on indirect evidence based on above‐ground plant responses. As a result, specific soil pathogen responses accompanying above‐ground plant community dynamics are largely unknown. Here, we examine the oomycete pathogens in soils conditioned by established populations of native noninvasive and non‐native invasive haplotypes of Phragmites australis (European common reed). Our aim was to assess whether populations of invasive plants harbor unique communities of pathogens that differ from those associated with noninvasive populations and whether the distribution of taxa within these communities may help to explain invasive success. We compared the composition and abundance of pathogenic and saprobic oomycete species over a 2‐year period. Despite a diversity of oomycete taxa detected in soils from both native and non‐native populations, pathogen communities from both invaded and noninvaded soils were dominated by species of Pythium. Pathogen species that contributed the most to the differences observed between invaded and noninvaded soils were distributed between invaded and noninvaded soils. However, the specific taxa in invaded soils responsible for community differences were distinct from those in noninvaded soils that contributed to community differences. Our results indicate that, despite the phylogenetic relatedness of native and non‐native P. australis haplotypes, pathogen communities associated with the dominant non‐native haplotype are distinct from those of the rare native haplotype. Pathogen taxa that dominate either noninvaded or invaded soils suggest different potential mechanisms of invasion facilitation. These findings are consistent with the hypothesis that non‐native plant species that dominate landscapes may “cultivate” a different soil pathogen community to their rhizosphere than those of rarer native species.     

Risk assessment model for the introduction of non‐native freshwater fish into New Zealand

A risk assessment model was developed to score the potential risk for both the establishment and impact of introduced freshwater fish species in New Zealand. Although based on similar models developed for Australia and the UK, it is customised to a New Zealand context in which the risk of a site‐specific ecological impact is of more concern than the potential for a species to spread rapidly and widely (i.e. its invasiveness). The model was calibrated using data on 21 introduced fish species already present in New Zealand and tested on eight species not present but for which a decision on introduction has been made using other methods. Threshold scores for risk of establishment, risk of impact and overall risk of causing ecological damage were set based on current knowledge and incorporated into a decision support system to provide managers with a numerical (vs subjective) basis for deciding on whether or not to permit the entry of a new species. The model assists in decision‐making on the introduction of new species and also provides a means of assessing the ecological risk posed by the further spread of those introduced fish already present.     

Laboratory and field validation of a simple method for detecting four species of non‐native freshwater fish using eDNA

This paper presents the first phase in the development and validation of a simple and reliable environmental (e)DNA method using conventional PCR to detect four species of non‐native freshwater fish: pumpkinseed Lepomis gibbosus, sunbleak Leucaspius delineatus, fathead minnow Pimephales promelas and topmouth gudgeon Pseudorasbora parva. The efficacy of the approach was demonstrated in indoor tank (44 l) trials in which all four species were detected within 24 h. Validation was through two field trials, in which L. gibbosus was detected 6–12 h after its introduction into outdoor experimental ponds and P. parva was successfully detected in disused fish rearing ponds where the species was known to exist. Thus, the filtration of small (30 ml) volumes of pond water was sufficient to capture fish eDNA and the approach emphasised the importance of taking multiple water samples of sufficient spatial coverage for detecting species of random or patchy distribution.     

Size structure of benthic freshwater fish communities in relation to environmental gradients

The benthic fish communities of 161 Swedish lakes, sampled with multi-mesh gillnets, were examined by direct gradient analyses with ten environmental factors. A geographic gradient, correlated with climatic factors and productivity, was most important for ordination of species optima. By contrast, the distribution of biomass in different size-classes and trophic groups, was influenced more directly by local factors such as water quality (pH) and lake morphometry (area and maximum depth). The communities are not only structured at the species level, but also by size-related allocation of resources, within and between coexisting species. This national survey confirmed patterns observed in previous local field studies, as well as experiments designed for testing mechanisms acting on size-structured populations. However, it highlighted the need for more extensive studies on if and how large-scale environmental variation affects the cues for size-related ontogenetic niche shifts in facultative piscivores, because species such as perch Perca fluviatilis , brown trout Salmo trutta , and Arctic charr Salvelinus alpinus often dominate the benthic fish community.