Spreading to a limit: the time required for a neophyte to reach its maximum range

The average range size of alien plants in Spain reaches a maximum at 143 years. This is consistent with estimates of such a maximum in Ireland, Britain, Germany and the Czech Republic. A round figure of about 150 years on average for neophytes to reach their maximum range in European countries is indicated.     

The distribution of range sizes of native and alien plants in four European countries and the effects of residence time

Aim Do the statistical distributions of range sizes of native and alien species differ? If so, is this because of residence time effects? And can such effects indicate an average time to a maximum? Location Ireland, Britain, Germany and the Czech Republic. Methods The data are presence or absence of higher plants in mapping units of 100 km² (Ireland and Britain) or c. 130 km² (Germany and the Czech Republic) in areas varying from 79 to 357 thousand km². Logit transforms of range sizes so defined were tested for normality, and examined by ANOVA, and by loess, ordinary least square (OLS) and reduced major axis regressions. Results Current range sizes, in logits, are near normally distributed. Those of native plants are larger than those of naturalized neophytes (plants introduced since 1500 ad ) and much larger than those of casual neophytes. Archaeophytes (introduced earlier) have range sizes slightly larger than natives, except in Ireland. Residence time, the time since an invasive species arrived in the wild at a certain place, affects range sizes. The relationships of the range of naturalized neophytes to residence time are effectively straight in all four places, showing no significant curvature or asymptote back to 1500, though there are few records between 1500 and 1800. The relationships have an r² of only about 10%. Both OLS regressions and reduced major axes can be used to estimate the time it takes for the range of a naturalized neophyte to reach a maximum. Main conclusions Established neophytes have smaller range size distributions than natives probably because many have not yet reached their maximum. We estimate it takes at least 150 years, possibly twice that, on average, for the maximum to be reached in areas of the order of 10⁵ km². Policy needs to allow for the variation in rates of spread and particularly the long time needed to fill ranges. Most naturalized neophytes are still expanding their ranges in Europe.     

Predicting and explaining plant invasions through analysis of source area floras: some critical considerations

Comparing species that become invasive with others from the same regional species pool that do not invade raises several issues about the accuracy of analyses attempting to define the determinants of invasiveness. The delimitation of the source area and deciding which species group(s) to include are especially relevant in analyses focusing on species originating in Europe. Historical patterns of immigration of alien species into Europe must be considered since European floras comprise a complex mix of native species, historical introductions (archaeophytes) and relative newcomers (neophytes). We make three main points: (1) Archaeophytes (species introduced to Europe before the discovery of America) differ from natives in a number of traits and in historical association with people; it is misleading to lump archaeophytes with native taxa. (2) Taxa from climatically and geographically different regions, representing distinct floristic geoelements, need to be treated separately, and not as a homogenous pool of potential invaders. Restricting the source species pool to native taxa with comparable phytogeographical characteristics reduces the variation associated with chance of dispersal by humans from the source area. (3) For prediction, a clear distinction should be made between accuracy (the proportion of those found to be alien that were also predicted to be there) and reliability (or predictive value, the proportion of those predicted to become aliens that do so). Information accumulated over centuries by botanists in Central Europe provides an excellent opportunity to deal with these issues and avoid spurious results. To illustrate these issues, we revisit a recently published study of Central‐European plant species as invaders in two Argentinean provinces ( Prinzing et al., 2002 ) to explore and demonstrate the implications of the above points. We hope that future studies will build on these points to achieve more reliable predictions.     

Assessing the potential range expansion of the exotic monk parakeet in Spain

In this study we determine favourable areas for the monk parakeet, Myiopsitta monachus , in peninsular Spain to account for its current distribution and predict its future course according to its potential range. We applied a favourability function based on generalized linear models using the presence/absence of breeding colonies of the species and the values of a set of variables on the 5167 UTM 10 × 10 km squares comprising the study area. We calculated the factor of distribution change in presences predicted by the model, and grouped the variables into explanatory factors performing a variation partitioning to assess the explanatory power of each factor. Our model included six predictors to explain the presence and absence of the species. These predictors were grouped into three factors: human activity, climate, and topography. Purely human influences accounted for 63.8% of the variation of the final model, while topographical variables explained 15.2% and climate only 5.7%. We obtained a high distribution change factor in which the presences of the species were predicted to increase between two- and sevenfold. Taking into account highly favourable squares, we conclude that the species is still absent in more than 72% of potential settlement areas, and thus we expect a continuous increase in the distribution of the species. Human activity is the main force moulding the distribution of the species, and lies behind its fast expansion, which is not only active, but is also passive via releases and escapes. We identified the areas of likely future expansion of the exotic monk parakeet in Spain. The pest status of the species in its native range, together to its distribution trend, should be taken into account by wildlife agencies to consider options for management.     

Controls on pathogen species richness in plants' introduced and native ranges: roles of residence time, range size and host traits

Introduced species escape many pathogens and other enemies, raising three questions. How quickly do introduced hosts accumulate pathogen species? What factors control pathogen species richness? Are these factors the same in the hosts' native and introduced ranges? We analysed fungal and viral pathogen species richness on 124 plant species in both their native European range and introduced North American range. Hosts introduced 400 years ago supported six times more pathogens than those introduced 40 years ago. In hosts' native range, pathogen richness was greater on hosts occurring in more habitat types, with a history of agricultural use and adapted to greater resource supplies. In hosts' introduced range, pathogen richness was correlated with host geographic range size, agricultural use and time since introduction, but not any measured biological traits. Introduced species have accumulated pathogens at rates that are slow relative to most ecological processes, and contingent on geographic and historic circumstance.     

The role of abiotic conditions in shaping the long-term patterns of a high-elevation Argentine ant invasion

Analysis of long‐term patterns of invasion can reveal the importance of abiotic factors in influencing invasion dynamics, and can help predict future patterns of spread. In the case of the invasive Argentine ant (Linepithema humile), most prior studies have investigated this species’ limitations in hot and dry climates. However, spatial and temporal patterns of spread involving two ant populations over the course of 30 years at a high elevation site in Hawaii suggest that cold and wet conditions have influenced both the ant's distribution and its rate of invasion. In Haleakala National Park on Maui, we found that a population invading at lower elevation is limited by increasing rainfall and presumably by associated decreasing temperatures. A second, higher elevation population has spread outward in all directions, but rates of spread in different directions appear to have been strongly influenced by differences in elevation and temperature. Patterns of foraging activity were strongly tied to soil temperatures, supporting the hypothesis that variation in temperature can influence rates of spread. Based on past patterns of spread, we predicted a total potential range that covers nearly 50% of the park and 75% of the park's subalpine habitats. We compared this rough estimate with point predictions derived from a degree‐day model for Argentine ant colony reproduction, and found that the two independent predictions match closely when soil temperatures are used in the model. The cold, wet conditions that have influenced Argentine ant invasion at this site are likely to be influential at other locations in this species’ current and future worldwide distribution.     

Tracking biological invasions in space and time: elucidating the invasive history of the green alga Codium fragile using old DNA

With the advent of 'ancient DNA' studies on preserved material of extant and extinct species, museums and herbaria now represent an important although still underutilized resource in molecular ecology. The ability to obtain sequence data from archived specimens can reveal the recent history of cryptic species and introductions. We have analysed extant and herbarium samples of the highly invasive green alga Codium fragile , many over 100 years old, to identify cryptic accessions of the invasive strain known as C. fragile ssp. tomentosoides , which can be identified by a unique haplotype. Molecular characterization of specimens previously identified as native in various regions shows that the invasive tomentosoides strain has been colonizing new habitats across the world for longer than records indicate, in some cases nearly 100 years before it was noticed. It can now be found in the ranges of all the other native haplotypes detected, several of which correspond to recognized subspecies. Within regions in the southern hemisphere there was a greater diversity of haplotypes than in the northern hemisphere, probably as a result of dispersal by the Antarctic Circumpolar Current. The findings of this study highlight the importance of herbaria in preserving contemporaneous records of invasions as they occur, especially when invasive taxa are cryptic.     

The times are changing: temporal shifts in patterns of fish invasions in central European fresh waters

This study examines the invasion history of alien fish species based on exhaustive national data sets on fish invasions of two contiguous central European countries (Germany and Austria). Fifteen alien fish species are currently established in both countries, constituting 14 and 17% of the total freshwater fish fauna of Germany and Austria, respectively. In both countries, six alien species are present, but not established. The status of five alien species in Germany and three species in Austria remains unknown. Accumulation rates of alien fish species have increased in recent decades with >50% of them reported after 1971. North America and Asia were the primary sources of alien fish species in Germany and Austria up to the 1980s, whereas European species of Ponto‐Caspian origin dominate now. Fisheries (including aquaculture) and the animal trade were responsible for most earlier introductions, whereas waterways were the main pathway for recent invaders. The extent of the spatial distribution of alien species was positively correlated with residence time, i.e. the time elapsed since the first national record. Different thermal preferences of early invaders (mostly coldwater species) and new invaders (typically warmwater adapted) may benefit the latter in the face of climate change. It is concluded that new challenges for alien fish management arise and that ecosystem‐based approaches as endorsed by the E.U. Water Framework Directive (maintaining or restoring good ecological status of rivers and streams) should become the centrepiece of river management in Europe.     

Native and naturalized range size in Pinus: relative importance of biogeography,introduction effort and species traits

Aim Pine trees (genus Pinus) represent an ancient lineage, naturally occurring almost exclusively in the Northern Hemisphere, but introduced and widely naturalized in both hemispheres. As large trees of interest to forestry, they attract much attention and their distribution is well documented in both indigenous and naturalized ranges. This creates an opportunity to analyse the relationship between indigenous and naturalized range sizes in the context of different levels of human usage, biological traits and the characteristics of the environments of origin. Location Global. Methods We combined and expanded pre‐existing data sets for pine species distributions and pine species traits, and used a variety of regression techniques (including generalized additive models and zero‐inflated Poisson models) to assess which variables explained naturalized and indigenous range sizes. Results Indigenous and naturalized range sizes are positively correlated but there are many notable exceptions. Some species have large indigenous ranges but small or no naturalized ranges, whereas others have small indigenous ranges, but have naturalized in many regions. Indigenous range is correlated to factors such as seed size (?), age at first reproduction (?), and latitude (+, supporting Rapoport's rule), but also to the extent of coverage of species in the forestry literature (+). Naturalized range size is strongly influenced by the extent of coverage of species in the forestry literature (+), a proxy for propagule pressure. Naturalization was also influenced by average elevation in the indigenous range (?) and age at first reproduction (?). Main conclusions The macroecological and evolutionary pressures facing plant groups are not directly transferable between indigenous and naturalized ranges. In particular, there are strong biases in species naturalization and expansion in invasive ranges that are unrelated to factors determining indigenous range size. At least for Pinus, a new set of macroecological patterns are emerging which are profoundly influenced by humans.     

Residence time,native range size,and genome size predict naturalization among angiosperms introduced to Australia

Although critical to progress in understanding (i) if, and (ii) at what rate, introduced plants will naturalize and potentially become invasive, establishing causal links between traits and invasion success is complicated by data gaps, phylogenetic nonindependence of species, the inability to control for differences between species in residence time and propagule pressure, and covariance among traits. Here, we focus on statistical relationships between genomic factors, life history traits, native range size, and naturalization status of angiosperms introduced to Australia. In a series of analyses, we alternately investigate the role of phylogeny, incorporate introduction history, and use graphical models to explore the network of conditional probabilities linking traits and introduction history to naturalization status. Applying this ensemble of methods to the largest publicly available data set on plant introductions and their fates, we found that, overall, residence time and native range size best predicted probability of naturalization. Yet, importantly, probability of naturalization consistently increased as genome size decreased, even when the effects of shared ancestry and residence time in Australia were accounted for, and that this pattern was stronger in species without a history of cultivation, but present across annual–biennials, and herbaceous and woody perennials. Thus, despite introduction biases and indirect effects of traits via introduction history, across analyses, reduced genome size was nevertheless consistently associated with a tendency to naturalize.     

Biological invasions drive size increases in marine and estuarine invertebrates

Ecologists have long been fascinated by the morphological changes that species frequently undergo when introduced into new regions. In this study an unusual pattern of size change associated with the invasion of 19 species of marine and estuarine invertebrates is reported. The results show that the majority of species are significantly larger in the introduced range compared with the native range with little evidence for any decrease in size following invasion. This invasion‐driven increase in body size sharply contrasts with the pattern observed in many other taxa including plants, mammals and lizards, where invaders frequently exhibit post‐invasion decreases and increases in size. These size changes were not influenced by differences in latitude, sample size or length of time since invasion. Although several mechanisms, may explain the results, none have been demonstrated.     

Exploring species attributes and site characteristics to assess plant invasions in Spain

Aim Biological invasions are a major component of global change with increasing effects on natural ecosystems and human societies. Here, we aim to assess the relationship between plant invader species attributes and the extent of their distribution range size, at the same time that we assess the association between environmental factors and plant invader species richness. Location Spain, Mediterranean region. Methods From the species perspective, we calculated the distribution range size of the 106 vascular plant invaders listed in a recently published atlas of alien plant species in Spain. Range size was used as an estimation of the degree of invasion success of the species. To model variation in range size between species as a function of a set of species attributes, we adopted the framework of the generalized linear mixed models because they allow the incorporation of taxonomic categories as nested random factors to control for phylogenetic relationships. From the invaded site perspective, we determined invader plant species richness as the number of species for each 10 × 10 km Universal Transverse Mercator (UTM) grid. For each grid cell, we estimated variables concerning landscape, topography, climate and human settlement. Then, we performed a generalized linear mixed model incorporating a defined spatial correlation structure to assess the relationship between plant invader richness and the environmental predictors. Results From the species perspective, wind dispersal and minimum residence time appeared to favour invasion success. From the invaded site perspective, we identified high anthropogenic disturbance, low altitude, short distance to the coastline and dry, hot weather as the main correlates to UTM grid cell invader richness. Main conclusions According to these results, an increasing importance of man‐modified ecosystems and global warming in the Mediterranean region should facilitate the expansion of plant invaders, especially wind‐dispersed species, leading to the accumulation of invasive species in some sites (i.e. invasion hot spots).     

Past and future of predictions in plant invasions: a field test by time

Intensive interest in the alien flora of the Czech Republic stimulated one of the earliest attempts (in the early 1970s) to predict potential invaders of arable land. Another paper published recently by the same research team using the same methodology provides a relatively unique possibility to assess the success of these predictions after a quarter of a century. The predictions were successful for 39.3% of the 28 species included, while 60.7% of invasion cases must be considered as failures. Prediction was rather unsuccessful for the members of Asteraceae (14.3%), and more correct for annuals/biennials (45.5%) than for perennials (16.7%). No pattern was found with respect to the area of origin. The results indicate that past predictions based largely on intuition were less successful than modern prediction systems using the knowledge of a large number of characters and carried out using advanced computation methods. The correct identification of invaders using such systems reaches values between 61% and 91%.     

Riparian vegetation: degradation, alien plant invasions, and restoration prospects

Rivers are conduits for materials and energy; this, the frequent and intense disturbances that these systems experience, and their narrow, linear nature, create problems for conservation of biodiversity and ecosystem functioning in the face of increasing human influence. In most parts of the world, riparian zones are highly modified. Changes caused by alien plants — or environmental changes that facilitate shifts in dominance creating novel ecosystems — are often important agents of perturbation in these systems. Many restoration projects are underway. Objective frameworks based on an understanding of biogeographical processes at different spatial scales (reach, segment, catchment), the specific relationships between invasive plants and resilience and ecosystem functioning, and realistic endpoints are needed to guide sustainable restoration initiatives. This paper examines the biogeography and the determinants of composition and structure of riparian vegetation in temperate and subtropical regions and conceptualizes the components of resilience in these systems. We consider changes to structure and functioning caused by, or associated with, alien plant invasions, in particular those that lead to breached abiotic‐ or biotic thresholds. These pose challenges when formulating restoration programmes. Pervasive and escalating human‐mediated changes to multiple factors and at a range of scales in riparian environments demand innovative and pragmatic approaches to restoration. The application of a new framework accommodating such complexity is demonstrated with reference to a hypothetical riparian ecosystem under three scenarios: (1) system unaffected by invasive plants; (2) system initially uninvaded, but with flood‐generated incursion of alien plants and escalating invasion‐driven alteration; and (3) system affected by both invasions and engineering interventions. The scheme has been used to derive a decision‐making framework for restoring riparian zones in South Africa and could guide similar initiatives in other parts of the world.     

Explaining invasiveness from the extent of native range: new insights from plant atlases and herbarium specimens

Aim

We tested the relationship between the extent of the native range and the success (number of occurrences) in the introduced range of European vascular plant species naturalized in the province of Québec (Canada). We hypothesized that the performance of models linking native range size and species invasiveness can be improved if residence time and climate tolerance are taken into account.

Methods

The extent of the native range (Europe, Asia) was estimated using plant atlases. The number of occurrences in the introduced range (Québec) was estimated using the number of herbarium specimens stored in herbaria. Herbarium specimens were also used to obtain residence time. Plant hardiness was used as an indicator of the suitability of a species to the climate of the introduced range. Multiple linear regression models, corrected to take into account phylogenetic biases, were used to calculate correlations between the extent of the native range and the number of occurrences in the introduced range.

Results

The larger the native distribution area in Eurasia, the greater the number of occurrences (herbarium specimens) in Québec. The shorter the residence time and the less hardy the plant, the fewer the number of occurrences. In all models tested, the phylogenetic structure explained a significant proportion of the variance, but its influence decreased as the number of species or area studied (Europe versus Eurasia) increased.

Main conclusions

The extent of the native range is a good explanatory variable for the invasion success of vascular plants, especially once other factors (residence time, climate tolerance, phylogeny) are taken into account. Thus, a model using these variables could be used by environmental managers to flag species warranting further investigation. With the emergence of online databases, gathering the required information is becoming easier and cheaper. As online databases continue to improve and new analytical tools are developed, this approach will become even more powerful.

     

Phenology in a warming world: differences between native and non‐native plant species

Phenology is a harbinger of climate change, with many species advancing flowering in response to rising temperatures. However, there is tremendous variation among species in phenological response to warming, and any phenological differences between native and non‐native species may influence invasion outcomes under global warming. We simulated global warming in the field and found that non‐native species flowered earlier and were more phenologically plastic to temperature than natives, which did not accelerate flowering in response to warming. Non‐native species' flowering also became more synchronous with other community members under warming. Earlier flowering was associated with greater geographic spread of non‐native species, implicating phenology as a potential trait associated with the successful establishment of non‐native species across large geographic regions. Such phenological differences in both timing and plasticity between native and non‐natives are hypothesised to promote invasion success and population persistence, potentially benefiting non‐native over native species under climate change.     

The vulnerability of habitats to plant invasion: disentangling the roles of propagule pressure,time and sampling effort

Aim To quantify the vulnerability of habitats to invasion by alien plants having accounted for the effects of propagule pressure, time and sampling effort. Location New Zealand. Methods We used spatial, temporal and habitat information taken from 9297 herbarium records of 301 alien plant species to examine the vulnerability of 11 terrestrial habitats to plant invasions. A null model that randomized species records across habitats was used to account for variation in sampling effort and to derive a relative measure of invasion based either on all records for a species or only its first record. The relative level of invasion was related to the average distance of each habitat from the nearest conurbation, which was used as a proxy for propagule pressure. The habitat in which a species was first recorded was compared to the habitats encountered for all records of that species to determine whether the initial habitat could predict subsequent habitat occupancy. Results Variation in sampling effort in space and time significantly masked the underlying vulnerability of habitats to plant invasions. Distance from the nearest conurbation had little effect on the relative level of invasion in each habitat, but the number of first records of each species significantly declined with increasing distance. While Urban, Streamside and Coastal habitats were over‐represented as sites of initial invasion, there was no evidence of major invasion hotspots from which alien plants might subsequently spread. Rather, the data suggest that certain habitats (especially Roadsides) readily accumulate alien plants from other habitats. Main conclusions Herbarium records combined with a suitable null model provide a powerful tool for assessing the relative vulnerability of habitats to plant invasion. The first records of alien plants tend to be found near conurbations, but this pattern disappears with subsequent spread. Regardless of the habitat where a species was first recorded, ultimately most alien plants spread to Roadside and Sparse habitats. This information suggests that such habitats may be useful targets for weed surveillance and monitoring.