Evaluating dominance as a component of non-native species invasions

Many studies have quantified plant invasions by determining patterns of non‐native species establishment (i.e. richness and absolute cover). Until recently, dominance has been largely overlooked as a significant component of invasion. Therefore, we re‐examined a 6‐year data set of 323 0.1 ha plots within 18 vegetation types collected in the Grand Staircase‐Escalante National Monument from 1998 to 2003, including dominance (i.e. relative cover) in our analyses. We specifically focused on the non‐native species Bromus tectorum, a notable dominant annual grass in this system. We found that non‐native species establishment and dominance are both occurring in species‐rich, mesic vegetation types. Therefore, non‐native species dominance may result despite many equally abundant native species rather than a dominant few, and competitive exclusion does not seem to be a primary control on either non‐native species establishment or dominance in this study. Unlike patterns observed for non‐native species establishment, relative non‐native species cover could not be predicted by native species richness across vegetation types (R² < 0.001; P = 0.45). However, non‐native species richness was found to be positively correlated with relative non‐native species cover and relative B. tectorum cover (R² = 0.46, P < 0.01; R² = 0.17, P < 0.01). Analyses within vegetation types revealed predominantly positive relationships among these variables for the correlations that were significant. Regression tree analyses across vegetation types that included additional biotic and abiotic variables were a little better at predicting non‐native species dominance (PRE = 0.49) and B. tectorum dominance (PRE = 0.39) than at predicting establishment. Land managers will need to set priorities for control efforts on the more productive, species‐rich vegetation types that appear to be susceptible to both components of invasion.     

The role of environmental gradients in non-native plant invasion into burnt areas of Yosemite National Park, California

Fire is known to facilitate the invasion of many non-native plant species, but how invasion into burnt areas varies along environmental gradients is not well-understood. We used two pre-existing data sets to analyse patterns of invasion by non-native plant species into burnt areas along gradients of topography, soil and vegetation structure in Yosemite National Park, California, USA. A total of 46 non-native species (all herbaceous) were recorded in the two data sets. They occurred in all seven of the major plant formations in the park, but were least common in subalpine and upper montane conifer forests. There was no significant difference in species richness or cover of non-natives between burnt and unburnt areas for either data set, and environmental gradients had a stronger effect on patterns of non-native species distribution, abundance and species composition than burning. Cover and species richness of non-natives had significant positive correlations with slope (steepness) and herbaceous cover, while species richness had significant negative correlations with elevation, the number of years post-burn, and cover of woody vegetation. Non-native species comprised a relatively minor component of the vegetation in both burnt and unburnt areas in Yosemite (percentage species = 4%, mean cover < 6.0%), and those species that did occur in burnt areas tended not to persist over time. The results indicate that in many western montane ecosystems, fire alone will not necessarily result in increased rates of invasion into burnt areas. However, it would be premature to conclude that non-native species could not affect post-fire succession patterns in these systems. Short fire-return intervals and high fire severity coupled with increased propagule pressure from areas used heavily by humans could still lead to high rates of invasion, establishment and spread even in highly protected areas such as Yosemite.     

Animal trade and non-indigenous species introduction: the world-wide spread of squirrels

Aim  In this study, a dataset on world-wide squirrel introductions has been used to locate the relative pathways and to determine the factors correlated with species establishment.
Location  The world.
Methods  The analysis includes a chronological table of introductions, a biogeographical analysis and an assessment of the likelihood of establishment according to species, propagule pressure, area of origin and characteristics of the recipient area.
Results  The main vector of such introductions was the intentional importation of live animals. Introductions increased in developed countries and proportionately to the volume of imported mammals. Moreover, areas characterized by higher numbers of native squirrels were more invaded. Squirrels were often introduced deliberately and only to a smaller extent escaped from captivity. The likelihood of their establishment increased proportionately to the number of animals released and decreased proportionately to the increase of the latitudinal distance between the recipient area and the native range of the species. The likelihood that the release of one pair of either Sciurus or Callosciurus species would establish a new population was higher than 50%.
Main conclusion  Squirrels proved to be successful invaders and their importation should be restricted so as to prevent further introductions.     

Sub-alpine amphibian distributions related to species palatability to non-native salmonids in the Klamath mountains of northern California

The goal of this study was to examine how introduced trout influence the distributions and abundances of a sub‐alpine amphibian assemblage whose members display a variety of different life‐history and defence strategies. Our study was conducted in the sub‐alpine lentic habitats of three wilderness areas that form the core of the Klamath‐Siskiyou Bioregion of northern California, a biodiversity ‘hotspot’ that supports the highest diversity of sub‐alpine, lentic‐breeding amphibians in the western USA. These wilderness areas contain no native fishes, but all have been populated with non‐native trout for recreational fishing. Five of the eight amphibian species that occur in this region were sufficiently common to use in our study; these included one that breeds in both temporary and permanent waters and is palatable to fish (Pacific treefrog, Pseudacris regilla), two that breed primarily in permanent waters and are unpalatable to fish (western toad, Bufo boreas, and rough‐skinned newt, Taricha granulosa), and two that breed primarily in permanent waters and are palatable to fish (Cascades frog, Rana cascadae, and long‐toed salamander, Ambystoma macrodactylum). Based on life histories and predator defence strategies (i.e. palatable or not), we predicted that the three palatable species would likely be negatively correlated with introduced trout, but with P. regilla less impacted because of its use of both temporary and permanent waters. We predicted that B. boreas and T. granulosa would not be significantly correlated with introduced trout due to the lack of any predator/prey interactions between them. We surveyed 728 pond, lake, or wet meadow sites during the summers of 1999–2002, using timed gill‐net sets to measure trout occurrence and relative density, and visual encounter surveys to determine amphibian presence and abundance. We used semiparametric logistic regression models to quantify the effect of trout presence/absence and density on the probability of finding amphibian species in a water body while accounting for variation within and among the various lentic habitats sampled. The distributions of P. regilla, A. macrodactylum and R. cascadae were strongly negatively correlated with trout presence across all three wilderness areas. Ambystoma macrodactylum was 44 times more likely to be found in lakes without fish than in lakes with fish. Rana cascadae and P. regilla were 3.7 and 3.0 times more likely, respectively, to be found in fishless than fish‐containing waters. In contrast, the two unpalatable species were either uncorrelated (T. granulosa) or positively correlated (B. boreas) with fish presence. We found that the relative density of fish (catch per unit effort) was negatively correlated with the combined abundances of the three palatable amphibians, and also with both the length and the condition of the fish themselves. Our results are consistent with a compelling body of evidence that introduced fishes greatly alter the aquatic community structure of mountain lakes, ponds, and wet meadows.     

Anthropogenic fires increase alien and native annual species in the Chilean coastal matorral

Aim We tested the hypothesis that anthropogenic fires favour the successful establishment of alien annual species to the detriment of natives in the Chilean coastal matorral. Location Valparaíso Region, central Chile. Methods We sampled seed rain, seedbank emergence and establishment of species in four paired burned and unburned areas and compared (using GLMM) fire resistance and propagule arrival of alien and native species. To assess the relative importance of seed dispersal and seedbank survival in explaining plant establishment after fire, we compared seed rain and seedbank structure with post‐fire vegetation using ordination analyses. Results Fire did not change the proportion of alien species in the coastal matorral. However, fire increased the number of annual species (natives and aliens) of which 87% were aliens. Fire reduced the alien seedbank and not the native seedbank, but alien species remained dominant in burned soil samples (66% of the total species richness). Seed rain was higher for alien annuals than for native annuals or perennials, thus contributing to their establishment after fire. Nevertheless, seed rain was less important than seedbank survival in explaining plant establishment in burned areas. Main conclusions Anthropogenic fires favoured alien and native annuals. Thus, fire did not increase the alien/native ratio but increased the richness of alien species. The successful establishment of alien annuals was attributable to their ability to maintain rich seedbanks in burned areas and to the greater propagule arrival compared to native species. The native seedbank also survived fire, indicating that the herbaceous community has become highly resilient after centuries of human disturbances. Our results demonstrate that fire is a relevant factor for the maintenance of alien‐dominated grasslands in the matorral and highlight the importance of considering the interactive effect of seed rain and seedbank survival to understand plant invasion patterns in fire‐prone ecosystems.     

Risk screening of non-native and translocated freshwater fish species in a Mediterranean-type shallow lake: Lake Marmara (West Anatolia)

Risk screening tools to identify species with a high or low risk of invasiveness are being increasingly used for effective management purposes. Amongst the available tools, the Fish Invasiveness Screening Kit (FISK) has been used extensively and successfully in large risk assessment (RA) areas, and was recently upgraded to the new generic tool Aquatic Species Invasiveness Screening Kit (AS-ISK). The aim of the present study was to assess with AS-ISK the invasive potential of introduced non-native and translocated fishes in a Mediterranean-type shallow lake (Lake Marmara) located in west Anatolia (Turkey). Based on independent evaluations of 35 species by two assessors, calibration of AS-ISK resulted in a threshold score of -3.65, which reliably distinguished between potentially invasive (high risk) and potentially non-invasive (medium to low risk) fishes. Of the 35 species assessed, 17 were categorised as ‘low risk’ and included native/endemic and translocated natives, and the remaining 18 as ‘high risk’ and comprised non-natives and translocated natives. Carassius gibelio had the highest score in the Climate Change Assessment section, suggesting that it might potentially impact on the native fish fauna under likely climate change scenarios for the RA area. Some cool water non-native (Oncorhynchus mykiss, Salvelinus fontinalis), translocated native (Luciobarbus lydianus) and endemic species (Ladigesocypris mermere) in the catchment will likely be affected negatively by predicted climate change conditions.     

Predicting the number of ecologically harmful exotic species in an aquatic system

Most introduced species apparently have little impact on native biodiversity, but the proliferation of human vectors that transport species worldwide increases the probability of a region being affected by high-impact invaders – i.e. those that cause severe declines in native species populations. Our study determined whether the number of high-impact invaders can be predicted from the total number of invaders in an area, after controlling for species–area effects. These two variables are positively correlated in a set of 16 invaded freshwater and marine systems from around the world. The relationship is a simple linear function; there is no evidence of synergistic or antagonistic effects of invaders across systems. A similar relationship is found for introduced freshwater fishes across 149 regions. In both data sets, high-impact invaders comprise approximately 10% of the total number of invaders. Although the mechanism driving this correlation is likely a sampling effect, it is not simply the proportional sampling of a constant number of repeat-offenders; in most cases, an invader is not reported to have strong impacts on native species in the majority of regions it invades. These findings link vector activity and the negative impacts of introduced species on biodiversity, and thus justify management efforts to reduce invasion rates even where numerous invasions have already occurred.     

Differential influence of Pomphorhynchus laevis (Acanthocephala) on the behaviour of native and invader gammarid species.

Although various species of acanthocephalan parasites can increase the vulnerability of their amphipod intermediate hosts to predation, particularly by altering their photophobic behaviour, their influence on the structure of amphipod communities and the success of invader species has so far received little attention. We compared the prevalence and behavioural influence of a fish acanthocephalan parasite, Pomphorhynchus laevis, in two species of amphipods, Gammarus pulex and Gammarus roeseli in sympatry in the river Ouche (Burgundy, eastern France). There, G. pulex is a resident species, whereas G. roeseli is a recent coloniser. Both uninfected G. pulex and G. roeseli were strongly photophobic, although less so in the invading species. However, there was no significant difference in reaction to light between infected and uninfected G. roeseli, whereas infected G. pulex were strongly photophilic. We discuss our results in relation to the parasite's ability to manipulate invading host species, the possibility that resistant individuals have been selected during the invasion process, and the role that acanthocephalan parasites can play in shaping the structure of amphipod communities.     

Global ecological impacts of invasive species in aquatic ecosystems

The introduction of invasive species, which often differ functionally from the components of the recipient community, generates ecological impacts that propagate along the food web. This review aims to determine how consistent the impacts of aquatic invasions are across taxa and habitats. To that end, we present a global meta‐analysis from 151 publications (733 cases), covering a wide range of invaders (primary producers, filter collectors, omnivores and predators), resident aquatic community components (macrophytes, phytoplankton, zooplankton, benthic invertebrates and fish) and habitats (rivers, lakes and estuaries). Our synthesis suggests a strong negative influence of invasive species on the abundance of aquatic communities, particularly macrophytes, zooplankton and fish. In contrast, there was no general evidence for a decrease in species diversity in invaded habitats, suggesting a time lag between rapid abundance changes and local extinctions. Invaded habitats showed increased water turbidity, nitrogen and organic matter concentration, which are related to the capacity of invaders to transform habitats and increase eutrophication. The expansion of invasive macrophytes caused the largest decrease in fish abundance, the filtering activity of filter collectors depleted planktonic communities, omnivores (including both facultative and obligate herbivores) were responsible for the greatest decline in macrophyte abundance, and benthic invertebrates were most negatively affected by the introduction of new predators. These impacts were relatively consistent across habitats and experimental approaches. Based on our results, we propose a framework of positive and negative links between invasive species at four trophic positions and the five different components of recipient communities. This framework incorporates both direct biotic interactions (predation, competition, grazing) and indirect changes to the water physicochemical conditions mediated by invaders (habitat alteration). Considering the strong trophic links that characterize aquatic ecosystems, this framework is relevant to anticipate the far‐reaching consequences of biological invasions on the structure and functionality of aquatic ecosystems.     

The impact of the invasive alien grass Cortaderia jubata (Lemoine) Stapf on an endangered mediterranean-type shrubland in California

Abstract. Since its introduction in the late 1800s, the perennial tussock grass Cortaderia jubata (Lemoine) Stapf has become an increasingly frequent member of coastal plant communities in California and Oregon. In this study, the community changes associated with C. jubata invasion into the mediterranean-type vegetation of Vandenberg Air Force Base, California were examined. Pristine plots of maritime chaparral were compared with spatially and topographically matched plots dominated by C. jubata . Aerial photographs indicated that the invaded plots had previously been shrubland. C. jubata invasion created a structurally less complex perennial grassland that was markedly depauperate in native shrub species. The absence of native shrubs depressed native richness in jubata grassland, but the greater richness of both native and alien herbaceous species made overall richness in jubata grassland indistinguishable from maritime chaparral. Vegetational differences were associated with differences in arthropod and small mammal populations between vegetation types. Arthropod abundance and order diversity were lower in plots dominated by C. jubata than in adjacent chaparral. Insect traps in C. jubata plots contained a significantly smaller proportion of Hymenoptera and Homoptera and a significantly greater proportion of Aràneae than traps in maritime chaparral. Rodent activity was significantly lower while rabbit activity was significantly greater in jubata grassland compared to maritime chaparral. This study indicates that the presence of C. jubata can dramatically alter the mediterranean-type landscape of central California. While it is likely that the initial establishment of C. jubata is associated with disturbances that are common in this ecosystem, the ability of C. jubata to expand from founding populations and to persist for long periods of time pose a serious threat to the native diversity of these unique systems.     

Vertebrate invaders and their ecological impacts in Chile

The South American country of Chile has been subjected to relatively few invasions in historic times. In comparison to the ca 610 vertebrate species found in continental Chile, only 24 species (3.9%) have invaded the country. Except for two bird and two or three mammal species that expanded their range and crossed the Andean Ranges from Argentina, the remaining 20 species were all introduced, or aided, by humans. Of the 24 invader species that currently occur in Chile, 11 (46%) were first introduced in central Chile, two (8%) in the northern part of the country, nine (38%) in the southern part, and two (8%) in Juan Fernández Archipelago. For ten (42%) of those species there is no information about their effects on the native flora, vegetation, or fauna. For eight species (33%), the information regarding the effects of the invaders is anecdotal. For six species (25%), there is adequate quantitative documentation available on their effects. Of a total of seven convincingly repo rted cases, two involve positive effects (increase in the prey base for native predators) and five have negative effects, of which three are on native vegetation and the remaining two on native fauna.     

Broad-scale determinants of non-native fish species richness are context-dependent

Identifying the factors determining the non-native species richness (NNSR) in a given area is essential for preventing species invasions. The relative importance of human-related and natural factors considered for explaining NNSR might depend upon both the spatial scale (i.e. the extent of the gradients sampled) and the historical context of the area surveyed. Here, using a worldwide database of freshwater fish occurrences, we tested whether the relative influence of human and ecological determinants of non-native fish species establishment at the scale of the biogeographic realm was consistent (i) with that observed worldwide, and (ii) among the different biogeographical realms. The prominent role of human activity in shaping the global (i.e. worldwide) pattern of NNSR cannot be directly extrapolated to the biogeographic realms. Furthermore, the relationships between human and ecological determinants and NNSR vary strikingly across biogeographic realms, revealing a strong context dependency of the determinants of NNSR. In particular, the human-related factors play a predominant role in explaining the establishment of non-native species in economically developed realms, while in the other realms environmental characteristics of the river basins best explained geographical patterns of NNSR. In the face of future biological invasions, considering both the spatial scale and the historical context of the surveyed area is crucial to adopt effective conservation strategies.     

Characteristics and habitats of non-native plant species in the city of Chonju,southern Korea

Investigations on non-native organisms have become an important task of modern ecology throughout the world. The major objective of this study was to identify the characteristics and habitats of non-native plant species in Korean cities in order to derive conclusions on the success of biological invaders. The city of Chonju in southwest Korea was selected as a representative investigation area. All wild-growing vascular plants were recorded in the city on a total of 106 sample plots located in representative land-use types and within various distances of the city center. Frequency, origin, life forms, and plant families of non-native species were analyzed. Additionally, the percentage of non-native species along a gradient from the city center to the outskirts was investigated. We found that most common non-native species of the flora of Chonju are: (i) native to the Northern hemisphere; (ii) belong to the annuals; (iii) belong to the plant families of the Compositae or Leguminosae; and (iv) they perform best on sites that are continuously disturbed by human impact. In conclusion, non-native species play a significant role in enhancing biological diversity in urban areas.     

外来种对生物多样性的影响及其控制

外来种是那些借助自身力量或其它外界力量传播到其未曾分布过的地域 ,并且能进行繁殖传播的生物。外来种入侵已成为一种引人关注的现象。外来种通过竞争、捕食、牧食、改变生境和传播疾病等方式对本地生物产生威胁 ,影响本地生物多样性。外来种入侵成了生物多样性丧失的两个主要影响之一 (另一影响是生境的破坏 )。因此 ,探寻阻止或减少外来种入侵的方法很有必要。目前 ,人们运用机械法、化学法和生物控制法来控制外来种。在外来种入侵的初始阶段或外来种数量不多时 ,运用机械法较好。化学法有带来新环境污染的危险。生物控制法是应用天敌来防治或消灭有害生物。在外来种的控制上 ,生物控制法有一些成功的案例 ,但它并不是万能的。有些生物控制剂可能对非目标种产生影响 ,这要求在释放前对生物控制剂作更严格的检测。     

Disturbance-driven changes in the variability of ecological patterns and processes

Understanding how disturbance shapes the dynamics of ecological systems is of fundamental importance in ecology. One emerging approach to revealing and appreciating disturbance effects involves examining disturbance-driven changes in the variability of ecological responses. Variability is rarely employed as a response variable to assess the influence of disturbance, but recent studies indicate that it can be an extremely sensitive metric, capturing differences obscured by averaging and conveying important ecological information about underlying causal processes. In this paper, we present a conceptual model to understand and predict the effects of disturbance on variability. The model estimates qualitative changes in variability by considering disturbance extent, frequency and intensity, as well as ecosystem recovery, and thereby captures not only the immediate effects of disturbance but also those that arise over time due to the biotic response to an event. We evaluate how well the model performs by comparing predictions with empirical results from studies examining a wide variety of disturbances and ecosystems, and discuss factors that may modify or even confound predictions. We include a concise guide to characterizing and detecting changes in variability, highlighting the most common and easily applied methods and conclude by describing several future directions for research. By considering variability as a response to disturbance, we gain another metric of fundamental system behaviour, an improved ability to identify organizing features of ecosystems and a better understanding of the predictability of disturbance-driven change – all critical aspects of assessing ecosystem response to disturbance.     

A multi-trait approach for the identification and protection of European freshwater species that are potentially vulnerable to the impacts of climate change

We present a multi-trait approach to identify potentially vulnerable species of Ephemeroptera (mayflies), Plecoptera (stoneflies) and Trichoptera (caddisflies), collectively referred to as EPT, to the impacts of climate change (CC). The “climate change vulnerability score” (CCVS) is an aggregation of six autecological traits that are known to be associated with vulnerability to CC: endemism, micro-endemism, temperature preference, altitudinal preference, stream zonation preference, and life history. We assigned a vulnerability score (0 – invulnerable to 6 – highly vulnerable to climate change) to 1940 EPT species and discussed the applicability of the index at three spatial scales: (1) continental (Europe), (2) state (the German Federal State of North Rhine-Westphalia) and (3) a river basin (the Ruhr River). We identified 157 EPT species (ca. 8%) as highly vulnerable to climate change (CCVS ≥ 4), including 95 species of caddisflies, 60 species of stoneflies and two species of mayflies. These are mostly found in France and Italy (52 species each), Spain and Slovenia (36 and 34, respectively), and Austria and Switzerland (30 species each), of which 95 are caddisflies, 60 stoneflies, and 2 mayflies. Using data collected in routine regional sampling we show that although no endemic EPTs were found in the German Federal State of North Rhine-Westphalia, eight species can still be identified as relatively vulnerable to CC (CCVS of 3). Almost all of these species are occurring in the ‘mountainous’ regions of the state (>200 m a.s.l.), the Sauerland and the Eifel. The upper reaches of the Ruhr catchment have been found to be relatively rich in vulnerable species, including several locally rare species. This index can assist conservationists to identify “hotspots” in terms of climate vulnerability and climate change refuge areas that can be considered for protection or the application of restoration measures at a local and regional scale. Nevertheless, not all species have complete autecological information, which hinders our ability to fully recognize the areas of priority. To further stabilize and enhance the applicability of this method, it is essential to fill these knowledge gaps in the future.     

The disruption of dominance hierarchies by a non-native species: an individual-based analysis

We studied the effects of the exotic rainbow trout (Oncorhynchus mykiss) on the performance and the dominance hierarchy of native Atlantic salmon (Salmo salar) at the group and individual level using laboratory and semi-natural experiments. At the group level, we compared the effects of interspecific and intraspecific competition (substitutive and additive design) on behavioural responses and growth of young-of-the-year Atlantic salmon. At the individual level, the same design was used to evaluate: (1) the temporal consistency of behavioural responses, dominance hierarchy and growth rate of Atlantic salmon; (2) the pattern of correlations between behaviours; and (3) the relationship between individual growth rate and behaviour. In the laboratory, group-level analyses revealed a weak but similar effect of rainbow trout and intraspecific competition on the behaviour and growth of Atlantic salmon. In contrast, individual-based analyses demonstrated that rainbow trout (but not intraspecific competition) strongly affected behavioural strategy, dominance hierarchy and growth trajectory of individual Atlantic salmon. Specifically, behaviours, dominance status and growth rate of salmon were temporally consistent in the intraspecific environment, while these patterns were disrupted when rainbow trout were present. Similarly, we found that rainbow trout strongly affected behavioural correlations and the relationships between individual growth rate and behaviour. The semi-natural experiments confirmed these results as interspecific competition affected relationships between individual growth rate of salmon, initial weight and activity index. Overall, individual-based analyses highlighted important mechanisms that were concealed at the group level, and that may be crucial to understand ecological and evolutionary consequences of exotic species. Moreover, these results demonstrated that competition with an exotic species disrupts the hierarchical relationship among native individuals and may therefore represent a potential for a shift in selective pressure.