The demography of introduction pathways,propagule pressure and occurrences of non‐native freshwater fish in England

• 1. Biological invasion theory predicts that the introduction and establishment of non‐native species is positively correlated with propagule pressure. Releases of pet and aquarium fishes to inland waters has a long history; however, few studies have examined the demographic basis of their importation and incidence in the wild.
• 2. For the 1500 grid squares (10×10 km) that make up England, data on human demographics (population density, numbers of pet shops, garden centres and fish farms), the numbers of non‐native freshwater fishes (from consented licences) imported in those grid squares (i.e. propagule pressure), and the reported incidences (in a national database) of non‐native fishes in the wild were used to examine spatial relationships between the occurrence of non‐native fishes and the demographic factors associated with propagule pressure, as well as to test whether the demographic factors are statistically reliable predictors of the incidence of non‐native fishes, and as such surrogate estimators of propagule pressure.
• 3. Principal coordinates of neighbour matrices analyses, used to generate spatially explicit models, and confirmatory factor analysis revealed that spatial distributions of non‐native species in England were significantly related to human population density, garden centre density and fish farm density. Human population density and the number of fish imports were identified as the best predictors of propagule pressure.
• 4. Human population density is an effective surrogate estimator of non‐native fish propagule pressure and can be used to predict likely areas of non‐native fish introductions. In conjunction with fish movements, where available, human population densities can be used to support biological invasion monitoring programmes across Europe (and perhaps globally) and to inform management decisions as regards the prioritization of areas for the control of non‐native fish introductions.

© Crown copyright 2010. Reproduced with the permission of her Majesty's Stationery Office. Published by John Wiley & Sons, Ltd.     

Metapopulation analysis indicates native and non‐native fishes respond differently to effects of wildfire on desert streams

Ash flows and flooding associated with wildfires represent important but understudied sources of disturbance for fish populations. Knowledge concerning these disturbances is especially limited for larger streams where warm water species dominate. Fire‐related disturbances have been hypothesised to differentially affect native and non‐native fishes, although this hypothesis has only been tested for salmonids. The objective of our research was to contrast effects of uncharacteristically large wildfires followed by flooding on metapopulations of native and non‐native fishes in the Gila River of southwest New Mexico. Probabilities of occupancy, colonisation and local extinction of fishes were calculated across sites before and during disturbance and were also measured across a broader spatial scale during disturbance to identify potential refuge locations. Occupancy was higher for native fishes than non‐natives, but multiple wildfire and flood events increased extinction probabilities of native species. Responses of non‐native species to wildfires were mixed; extinction of non‐native salmonids increased during disturbance, while extinction of several warm water species remained unchanged or decreased. Several undisturbed sites were poor refugia for natives as they were impacted by non‐native piscivores, dewatering, and fragmentation. However, despite exposure to multiple disturbances, sites located in large tributary and valley reaches were consistently occupied by native species, suggesting these habitats provided refugia. We suggest that management actions (forest thinning; prescribed burning) that restore a more natural disturbance regime of small and less severe fires coupled with habitat remediation activities (non‐native removal; decreased water withdrawal; improved connectivity) might diminish extinction risk for native fishes exposed to wildfire disturbance.     

Occupancy dynamics of rare cyprinids after invasive fish eradication

1. The eradication of invasive species is a management strategy implemented to eliminate adverse impacts of invaders on native species communities. After eradications, follow‐up studies are done to confirm eradication of the invasive species and the short‐term recovery of the native species, but long‐term monitoring to confirm full population recovery and stability is often not completed.
2. In this study, long‐term monitoring of native fishes was carried out over 5 years after the eradication of an invasive fish from the Rondegat River, South Africa. Forty‐six sites distributed along four river sections were sampled for presence/absence using underwater cameras and snorkel surveys. Density data were collected by snorkel surveys.
3. Using multi‐season occupancy models, the annual probability of colonization and local extinction of the native fishes and annual rate of change in occupancy along the river were estimated. Changes in native fish densities across time and across the control and treatment sections were analysed using Kruskal–Wallis analysis, followed by Dunn's post‐hoc test.
4. Probability of colonization and local extinction differed for each native fish species and may have been affected by extrinsic factors, such as rainfall, and intrinsic density‐dependent factors, hypothesized from the density data. The occupancy rates of change revealed that the two Near Threatened fish species have reached an occupancy dynamic equilibrium but the Endangered fish has not, suggesting that other conservation efforts may be needed.
5. Long‐term monitoring of native fishes after an eradication programme has confirmed the successful removal of the invader and the recovery and stability of the community. However, successful eradication was not sufficient for full recovery of all species; additional conservation management strategies are needed to secure the population stability and persistence of endangered fishes. We recommend that eradication programmes, regardless of locality, should employ long‐term monitoring to ensure full recovery of a native fish community.

     

Scientific uncertainty and the assessment of risks posed by non-native freshwater fishes

The introduction of non‐indigenous plants, animals and pathogens is a pressing global environmental challenge. Although not all introduced species become established and the fraction of those that do often have little appreciable effect on their new ecosystems, many others exert significant ecological, evolutionary and economic impacts. Stimulating further debate, Gozlan [Fish and Fisheries (2008) Vol. 9, pp. 106–115] argued that the majority of intentional freshwater fish introductions associated with aquaculture (fish species providing societal benefits) have not been reported as having an ecological impact. We find little to argue with his suggestion that low risk of ecological impact coupled with high market value encourages further introductions. But do we have an adequate understanding of the ecological risks associated with fish introductions to support such decisions? Indeed, resource managers and decision makers require some scientific knowledge to support their management actions; without this information, a precautionary approach is the only sensible course of action. The precautionary approach implies that the lack of scientific certainty is reason enough for postponing intentional introduction of non‐native species to avoid potentially serious or irreversible harm to the environment. Here, we suggest that we actually know very little about ecological impacts associated with fish introductions and that it would be therefore wholly inappropriate to equate a lack of data with a conclusion of ‘no impact’. We discuss four major challenges for enhancing the assessment of risks posed by non‐native freshwater fishes in the face of scientific uncertainty and highlight research opportunities and some alternative approaches for confronting these challenges in the future.     

Longer food chains and crowded niche space: effects of multiple invaders on desert stream food web structure

Tributaries of the Colorado River Basin, historically home to a complex of endemic omnivores collectively referred to as the ‘three species’; flannelmouth sucker (Catostomus latipinnis), bluehead sucker (C. discobolus) and roundtail chub (Gila robusta), have experienced the establishment of numerous non‐native fish species. In this study, we examine the impacts of the trophic ecology of non‐native fishes on the ‘three species’ in the San Rafael River, Utah, USA. We employ a suite of abundance comparisons, stable isotope techniques and size‐at‐age back‐calculation analyses to compare food web structure and growth rates of the ‘three species’ in study areas with and without established populations of non‐native species. We found that the ‘three species’ are more abundant in areas with few non‐native fishes present, regardless of habitat complexity. Stable isotope analyses indicate non‐native fishes lengthen the food chain by 0.5 trophic positions. Further, the trophic niche spaces of the native fishes shift and are narrower in the presence of non‐native fishes, as several non‐native species’ trophic niche spaces overlap almost entirely with each of the ‘three species’ (bluehead sucker and flannelmouth sucker 100%, roundtail chub 98.5%) indicating strong potential for competition. However, the ‘three species’ demonstrated no evidence of reduced growth in the presence of these non‐native fishes. Collectively, these results suggest that while non‐native fishes alter the food web structure presenting novel sources of predation and competition, mechanisms other than competition are controlling the size‐structure of ‘three species’ populations in the San Rafael River.     

Beaver dams shift desert fish assemblages toward dominance by non‐native species (Verde River,Arizona, USA)

The reintroduction of beaver (Castor canadensis) into arid and semi‐arid rivers is receiving increasing management and conservation attention in recent years, yet very little is known about native versus non‐native fish occupancy in beaver pond habitats. Streams of the American Southwest support a highly endemic, highly endangered native fish fauna and abundant non‐native fishes, and here we investigated the hypothesis that beaver ponds in this region may lead to fish assemblages dominated by non‐native species that favour slower‐water habitat. We sampled fish assemblages within beaver ponds and within unimpounded lotic stream reaches in the mainstem and in tributaries of the free‐flowing upper Verde River, Arizona, USA. Non‐native fishes consistently outnumbered native species, and this dominance was greater in pond than in lotic assemblages. Few native species were recorded within ponds. Multivariate analysis indicated that fish assemblages in beaver ponds were distinct from those in lotic reaches, in both mainstem and tributary locations. Individual species driving this distinction included abundant non‐native green sunfish (Lepomis cyanellus) and western mosquitofish (Gambusia affinis) in pond sites, and native desert sucker (Catostomus clarkii) in lotic sites. Overall, this study provides the first evidence that, relative to unimpounded lotic habitat, beaver ponds in arid and semi‐arid rivers support abundant non‐native fishes; these ponds could thus serve as important non‐native source areas and negatively impact co‐occurring native fish populations.     

Effect of an intensive mechanical removal effort on a population of non‐native rainbow trout Oncorhynchus mykiss in a South African headwater stream

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Vulnerability of Cape Fold Ecoregion freshwater fishes to climate change and other human impacts

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Can translocated native fishes retain their trophic niche when confronted with a resident invasive?

Diet interactions between native and non‐native fishes may influence the establishment of native species within their historical range (i.e., reintroduction). Therefore, we illustrated the food web structure of and followed the transition of the federally endangered humpback chub Gila cypha into a novel food web following translocation and determined the potential for a non‐native species, rainbow trout Oncorhynchus mykiss, to influence translocation success. Humpback chub and rainbow trout used resources high in the food web and assimilated similar proportions of native fishes, suggesting non‐native rainbow trout may occupy an ecological role similar to humpback chub. Subsequently, humpback chub may be well suited to colonise tributaries because of their ability to consume resources high in the food web. Additionally, diet partitioning may occur between all members of the fish community as indicated by separation in trophic niche space and little trophic overlap; although all species, particularly bluehead sucker Catostomus discobolus, used a broad range of food resources. Rainbow trout stomach content analysis corroborated stable isotope analysis and suggested rainbow trout diet consisted of aquatic and terrestrial macroinvertebrates, while larger rainbow trout (>120 mm total length) consumed a greater proportion of fish (incidence of piscivory = 5.3%). Trophic interactions may reveal an underutilized niche space or biotic resistance to the establishment of translocated native fishes. Continued translocation of humpback chub into tributaries appears to be one option for conservation. However, successful establishment of humpback chub may depend on continued removal of non‐native trout, increasing availability of diet sources at higher trophic levels.     

Spatial distribution patterns of the non‐native European catfish,Silurus glanis,from multiple online sources – a case study for the River Tagus (Iberian Peninsula)

Effective management of invasive fishes depends on the availability of updated information about their distribution and spatial dispersion. Forensic analysis was performed using online and published data on the European catfish, Silurus glanis L., a recent invader in the Tagus catchment (Iberian Peninsula). Eighty records were obtained mainly from anglers’ fora and blogs, and more recently from www.youtube.com . Since the first record in 1998, S. glanis expanded its geographic range by ≈700 km of river network, occurring mainly in reservoirs and in high‐order reaches. Human‐mediated and natural dispersal events were identified, with the former occurring during the first years of invasion and involving movements of >50 km. Downstream dispersal directionality was predominant. The analysis of online data from anglers was found to provide useful information on the distribution and dispersal patterns of this non‐native fish, and is potentially applicable as a preliminary, exploratory assessment tool for other non‐native fishes.     

Environmental DNA as a non‐invasive sampling tool to detect the spawning distribution of European anadromous shads (Alosa spp.)

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Diet of non‐native northern snakehead (Channa argus) compared to three co‐occurring predators in the lower Potomac River,USA

Abstract – Introductions of large, non‐native, carnivorous fishes continue to occur worldwide and represent a substantial management concern to global biodiversity. One of the most recent non‐native fishes to successfully establish in North America is the northern snakehead (Channa argus), found in the lower Potomac River catchment. Dispersal of the northern snakehead throughout this system has been well documented since its original discovery in May 2004; however, little is known about the foraging habits of this species and its interactions with co‐occurring predators. Here, we quantify northern snakehead diet in comparison with the diets of naturalised largemouth bass (Micropterus salmoides), and native American eel (Anguilla rostrata) and yellow perch (Perca flavescens) collected from tidal freshwaters bordering Virginia and Maryland near Fort Belvoir, Virginia. Over 97% of northern snakehead gut contents were fishes, with fundulid and centrarchid species consumed most frequently. Dietary overlap was biologically significant only between northern snakehead and largemouth bass. Aquatic invertebrates were >10 times more common in native predator diets, reducing dietary overlap with northern snakehead. Ontogenic shifts in adult northern snakehead diet were also detected, which may be explained by optimal foraging rather than true prey specificity. Northern snakehead may be occupying a novel niche based on a piscivorous diet, therefore limiting competition with resident predators in the lower Potomac River. Further research into interactions between largemouth bass and northern snakehead is needed to inform management decisions and understand the ecological impacts of this non‐native species.     

Nile tilapia (Oreochromis niloticus) invasions disrupt the functional patterns of fish community in a large subtropical river in China

Although non‐native species can modify ecosystem function, their effects in southern China have not been investigated, despite their ecological importance. This study assessed how Nile tilapia Oreochromis niloticus (L.) invasions impact the functional patterns of fish communities in the large subtropical Pearl River by analysing the relationships between invasion and body size over a 9‐year period. The relative abundance of Nile tilapia increased significantly over time. In particular, the established Nile tilapia significantly undermined the relative densities and the body size of the native fishes. Importantly, this study provided empirical evidence that functional indices (such as body size) are more sensitive when evaluating the effects of non‐native species on a community than richness. Understanding processes such as those should be the basis of controlling alien fish species and fisheries management in the Pearl River.     

Non‐native marine species in north‐west Europe: Developing an approach to assess future spread using regional downscaled climate projections

1. Climate change can affect the survival, colonization and establishment of non‐native species. Many non‐native species common in Europe are spreading northwards as seawater temperatures increase. The similarity of climatic conditions between source and recipient areas is assumed to influence the establishment of such species, however, in a changing climate those conditions are difficult to predict. 2. A risk assessment methodology has been applied to identify non‐native species with proven invasive qualities that have not yet arrived in north‐west Europe, but which could become problematic in the future. Those species with the highest potential to become established or be problematic have been taken forward, as well as some that may be economically beneficial, for species distribution modelling to determine future potential habitat distributions under projected climate change. 3. In the past, species distribution models have usually made use of low resolution global environmental datasets. Here, to increase the local resolution of the distribution models, downscaled shelf seas climate change model outputs for north‐west Europe were nested within global outputs. In this way the distribution model could be trained using the global species presence data including the species' native locations, and then projected using more comprehensive shelf seas data to understand habitat suitability in a potential recipient area. 4. Distribution modelling found that habitat suitability will generally increase further north for those species with the highest potential to become established or problematic. Most of these are known to be species with potentially serious consequences for conservation. With caution, a small number of species may present an opportunity for the fishing industry or aquaculture. The ability to provide potential future distributions could be valuable in prioritizing species for monitoring or eradication programmes, increasing the chances of identifying problem species early. This is particularly important for vulnerable infrastructure or protected or threatened ecosystems.     

Pan‐continental invasion of Pseudorasbora parva: towards a better understanding of freshwater fish invasions

In recent years, policy‐makers have sought the development of appropriate tools to prevent and manage introductions of invasive species. However, these tools are not well suited for introductions of non‐target species that are unknowingly released alongside intentionally‐introduced species. The most compelling example of such invasion is arguably the topmouth gudgeon Pseudorasbora parva, a small cyprinid species originating from East Asia. A combination of sociological, economical and biological factors has fuelled their rapid invasion since the 1960s; 32 countries (from Central Asia to North Africa) have been invaded in less than 50 years. Based on a combination of monitoring surveys (2535 populations sampled) and literature reviews, this paper aims to quantify and characterise important invasion parameters, such as pathways of introduction, time between introduction and detection, lag phase and plasticity of life history traits. Every decade, five new countries have reported P. parva introduction, mainly resulting from the movement of Chinese carps for fish farming. The mean detection period after first introduction was 4 years, a duration insufficient to prevent their pan‐continental invasion. High phenotypic plasticity in fitness related traits such as growth, early maturity, fecundity, reproductive behaviour and the ability to cope with novel pathogens has predisposed P. parva to being a strong invader. The Pseudorasbora parva invasion has provided quantitative data for the development of 1) early warning systems across different spatial scales; 2) rapid eradication programmes prior to natural spread in open systems and 3) sound risk assessments with emphasis on plasticity of life history traits.     

Risk screening of non‐native freshwater fishes in Croatia and Slovenia using the Fish Invasiveness Screening Kit

The Fish Invasiveness Screening Kit (FISK) version 2 was used to assess the invasiveness potential of 40 introduced and translocated freshwater fish species to Croatia and Slovenia. Based on a priori classification of invasiveness, receiver operating characteristic analysis of FISK scores from two independent assessors resulted in a statistically significant calibration threshold of 11.75. This indicated that FISK was able to discriminate reliably between non‐native species likely to pose a high risk of being invasive and those likely to pose a medium or low risk of invasiveness. Seven species were categorised as ‘medium risk’ and the other 33 as ‘high risk’, whereas no species was categorised as ‘low risk’. The two highest scoring species were European catfish Silurus glanis and North African catfish Clarias gariepinus. Mean scores for all species classified a priori as invasive were ranked as ‘high risk’ sensu lato and fell into the ‘moderately high risk’ subcategory. FISK proved to be a valid tool for assessing the risks posed by non‐native fishes in Croatia and Slovenia. For this reason, it can be adopted as a reliable tool for the prevention of new translocations or introductions of potentially invasive species in the risk assessment area, as well as to assist in decisions regarding future management (i.e. monitoring, control and eradication) and conservation strategies.     

Long‐term changes of marine subtidal benthic communities in North East Asia (Yellow and Japan seas) in a global change context: A review

1. A review of the long‐term changes and variations in benthic communities and the current status of the marine invasive species (MIS) in shallow waters of the Yellow Sea (Chinese sector) and the Sea of Japan (Russian and partly Korean sectors) is presented. This paper reflects on the progress and lessons learned, recommending actions for the future about the conservation of biodiversity.
2. In the Bohai Sea, the benthic ecosystem has been degenerating due to anthropogenic activities such as overfishing and pollution since the 1950s. The dominant position of K‐strategy species is gradually being lost and replaced by R‐strategy species. In the Yellow Sea, the macrobenthic community is different from other areas due to the Yellow Sea Cold Water Mass. Many economic species have been destroyed, and the biotic structure has changed significantly due to overfishing and climate change.
3. In the Russian sector of the Sea of Japan, the macrobenthic communities in the shallow‐water soft bottom have generally been in a stable condition for the last decades, except for some heavily polluted or disturbed areas due to dredging operations. The abundance of select large invertebrate species has changed considerably due to commercial fishing and poaching. Variations in macro? and meiobenthic communities under aquaculture conditions have occurred on a local scale during the last five decades.
4. MIS show obvious differences between China and Russia in the following aspects: introduction pathways of MIS, composition and number of non‐native species, threats and impacts of MIS to native communities and ecosystems, and economic and public health impacts.
5. Long‐term monitoring programmes should be developed to reveal future biotic changes and to separate the effects of cyclic variations of benthic communities from the impacts of pollution and eutrophication. Standardization of sampling procedures is required to compare changes/alternations in benthos across various regions worldwide.

     

Parasitism of a native Hawaiian stream fish by an introduced nematode increases with declining precipitation across a natural rainfall gradient

Climate change can promote disease emergence if shifting conditions favour infection of native fauna by introduced parasites and pathogens. In Hawai'i, climate warming is predicted to reduce net precipitation and surface flow in streams, which in turn could increase parasitism of native stream fishes by non‐native parasites. In this study, we utilised a natural precipitation gradient across the Hamakua coast on the island of Hawai'i to assess the relationship between precipitation and infection of Awaous stamineus, a native amphidromous goby, by the introduced nematode Camallanus cotti. We found that the abundance, intensity and prevalence of C. cotti in A. stamineus increased with declining rainfall. Our results also show that parasitism tracks precipitation patterns across the archipelago and that parasitism increases even with moderate decreases in rainfall. As the Hamakua coast precipitation gradient represents a proxy for predicted climate‐driven reductions in precipitation, these findings suggest that infection of native Hawaiian fishes by introduced parasites will increase if climate conditions change as expected. Our findings also suggest that parasitism may be exacerbated by other factors that reduce surface flow, including water extraction for agricultural and urban uses. If so, then adaptive management of minimum flow standards in Hawai'i and elsewhere could improve the well‐being of at‐risk native fishes by alleviating parasitism under current and future climate conditions.     

Voracious invader or benign feline? A review of the environmental biology of European catfish Silurus glanis in its native and introduced ranges*

A popular species for food and sport, the European catfish (Silurus glanis) is well‐studied in its native range, but little studied in its introduced range. Silurus glanis is the largest‐bodied freshwater fish of Europe and is historically known to take a wide range of food items including human remains. As a result of its piscivorous diet, S. glanis is assumed to be an invasive fish species presenting a risk to native species and ecosystems. To assess the potential risks of S. glanis introductions, published and ‘grey’ literature on the species’ environmental biology (but not aquaculture) was extensively reviewed. Silurus glanis appears well adapted to, and sufficiently robust for, translocation and introduction outside its native range. A nest‐guarding species, S. glanis is long‐lived, rather sedentary and produces relatively fewer eggs per body mass than many fish species. It appears to establish relatively easily, although more so in warmer (i.e. Mediterranean) than in northern countries (e.g. Belgium, UK). Telemetry data suggest that dispersal is linked to flooding/spates and human translation of the species. Potential impacts in its introduced European range include disease transmission, hybridization (in Greece with native endemic Aristotle’s catfish [Silurus aristotelis]), predation on native species and possibly the modification of food web structure in some regions. However, S. glanis has also been reported (France, Spain, Turkmenistan) to prey intensively on other non‐native species and in its native Germany to be a poor biomanipulation tool for top‐down predation of zooplanktivorous fishes. As such, S. glanis is unlikely to exert trophic pressure on native fishes except in circumstances where other human impacts are already in force. In summary, virtually all aspects of the environmental biology of introduced S. glanis require further study to determine the potential risks of its introduction to novel environments.     

Reconstruction of the historical distribution ranges of imperilled stream fishes from a global endemic hotspot based on molecular data: Implications for conservation of threatened taxa

1. Understanding historical distribution patterns of freshwater fishes prior to human impacts is crucial for informing effective strategies for biodiversity conservation. However, incomplete information on species occurrence records, the existence of cryptic species and sensitivity to small sample sizes limit the application of historical records in natural history collections as well as conventional species distribution modelling algorithms to infer past distributions of species.
2. This study used molecular data as an alternative and objective approach to reconstruct the historical distribution ranges of four stream fishes from the Breede River system in the Cape Fold Ecoregion, a global hotspot of imperilled endemic freshwater biodiversity in southern Africa.
3. The study used 249 occurrence records and 208 mitochondrial cytochrome b sequences to reconstruct the potential historical ranges of four taxa: Galaxias sp. ‘zebratus nebula', Galaxias sp. ‘zebratus Riviersonderend', Pseudobarbus sp. ‘burchelli Breede' and Pseudobarbus skeltoni.
4. All four taxa historically had broader distribution ranges across the Breede River system before human impacts, but they have suffered severe attrition as the main‐stem populations have been extirpated.
5. The severe decline in the historical ranges of these four taxa is a result of multiple impacts, particularly hydrological modification, habitat degradation and the introduction of non‐native species, which are also global challenges for freshwater ecosystems.
6. The approach presented in this study has great potential for reconstructing historical ranges of stream‐dwelling taxa from disparate regions where fragmentation has resulted from human‐mediated impacts. This information is crucial for identifying appropriate conservation strategies such as river rehabilitation and eradication of non‐native species, as well as guiding reintroductions and informing assisted gene flow where these are deemed necessary interventions.