Patterns of plant naturalization show that facultative mycorrhizal plants are more likely to succeed outside their native Eurasian ranges

The naturalization of an introduced species is a key stage during the invasion process. Therefore, identifying the traits that favor the naturalization of non-native species can help understand why some species are more successful when introduced to new regions. The ability and the requirement of a plant species to form a mutualism with mycorrhizal fungi, together with the types of associations formed may play a central role in the naturalization success of different plant species. To test the relationship between plant naturalization success and their mycorrhizal associations we analysed a database composed of mycorrhizal status and type for 1981 species, covering 155 families and 822 genera of plants from Europe and Asia, and matched it with the most comprehensive database of naturalized alien species across the world (GloNAF). In mainland regions, we found that the number of naturalized regions was highest for facultative mycorrhizal, followed by obligate mycorrhizal and lowest for non-mycorrhizal plants, suggesting that the ability of forming mycorrhizas is an advantage for introduced plants. We considered the following mycorrhizal types: arbuscular, ectomycorrhizal, ericoid and orchid mycorrhizal plants. Further, dual mycorrhizal species were those that included observations of arbuscular mycorrhizas as well as observations of ectomycorrhizas. Naturalization success (based on the number of naturalized regions) was highest for arbuscular mycorrhizal and dual mycorrhizal plants, which may be related to the low host specificity of arbuscular mycorrhizal fungi and the consequent high availability of arbuscular mycorrhizal fungal partners. However, these patterns of naturalization success were erased in islands, suggesting that the ability to form mycorrhizas may not be an advantage for establishing self-sustaining populations in isolated regions. Taken together our results show that mycorrhizal status and type play a central role in the naturalization process of introduced plants in many regions, but that their effect is modulated by other factors.     

Effects of climate change and horticultural use on the spread of naturalized alien garden plants in Europe

Climate warming is supposed to enlarge the area climatically suitable to the naturalization of alien garden plants in temperate regions. However, the effects of a changing climate on the spread of naturalized ornamentals have not been evaluated by spatially and temporarily explicit range modelling at larger scales so far. Here, we assess how climate change and the frequency of cultivation interactively determine the spread of 15 ornamental plants over the 21st century in Europe. We coupled species distribution modelling with simulations of demography and dispersal to predict range dynamics of these species in annual steps across a 250 × 250 m raster of the study area. Models were run under four scenarios of climate warming and six levels of cultivation intensity. Cultivation frequency was implemented as size of the area used for planting a species. Although the area climatically suitable to the 15 species increases, on average, the area predicted to be occupied by them in 2090 shrinks under two of the three climate change scenarios. This contradiction obviously arises from dispersal limitations that were pronounced although we assumed that cultivation is spatially adapting to the changing climate. Cultivation frequency had a much stronger effect on species spread than climate change, and this effect was non‐linear. The area occupied increased sharply from low to moderate levels of cultivation intensity, but levelled off afterwards. Our simulations suggest that climate warming will not necessarily foster the spread of alien garden plants in Europe over the next decades. However, climatically suitable areas do increase and hence an invasion debt is likely accumulating. Restricting cultivation of species can be effective in preventing species spread, irrespective of how the climate develops. However, for being successful, they depend on high levels of compliance to keep propagule pressure at a low level.     

Traits explain invasion of alien plants into tropical rainforests

• 1. The establishment of new botanic gardens in tropical regions highlights a need for weed risk assessment tools suitable for tropical ecosystems. The relevance of plant traits for invasion into tropical rainforests has not been well studied.
• 2. Working in and around four botanic gardens in Indonesia where 590 alien species have been planted, we estimated the effect of four plant traits, plus time since species introduction, on: (a) the naturalization probability and (b) abundance (density) of naturalized species in adjacent native tropical rainforests; and (c) the distance that naturalized alien plants have spread from the botanic gardens.
• 3. We found that specific leaf area (SLA) strongly differentiated 23 naturalized from 78 non‐naturalized alien species (randomly selected from 577 non‐naturalized species) in our study. These trends may indicate that aliens with high SLA, which had a higher probability of naturalization, benefit from at least two factors when establishing in tropical forests: high growth rates and occupation of forest gaps. Naturalized aliens had high SLA and tended to be short. However, plant height was not significantly related to species'' naturalization probability when considered alongside other traits.
• 4. Alien species that were present in the gardens for over 30 years and those with small seeds also had higher probabilities of becoming naturalized, indicating that garden plants can invade the understorey of closed canopy tropical rainforests, especially when invading species are shade tolerant and have sufficient time to establish.
• 5. On average, alien species that were not animal dispersed spread 78 m further into the forests and were more likely to naturalize than animal‐dispersed species. We did not detect relationships between the measured traits and estimated density of naturalized aliens in the adjacent forests.
• 6. Synthesis: Traits were able to differentiate alien species from botanic gardens that naturalized in native forest from those that did not; this is promising for developing trait‐based risk assessment in the tropics. To limit the risk of invasion and spread into adjacent native forests, we suggest tropical botanic gardens avoid planting alien species with fast carbon capture strategies and those that are shade tolerant.

     

Plant introduction,naturalization, and invasion in French Guiana (South America)

Continental tropical ecosystems are generally viewed as less vulnerable to biological invasions than island ones. Their apparent resistance to invasive alien species is often attributed to their higher native biota diversity and complexity. However, with the increase of human activities and disturbances and the accelerate rate of introductions of plant species, these apparently resilient continental ecosystems are now experiencing alien plant naturalization and invasion events. In order to illustrate this emergent phenomenon, we compiled a list of all known introduced and naturalized plant species in French Guiana (Guiana Shield, South America). A total of 490 alien plants were recorded, about 34% of which are currently naturalized, mainly species belonging to the Acanthaceae and Fabaceae (Faboideae) in the Eudicotyledons, and Poaceae (grasses) and Arecaceae (palms) in the Monocotyledons. The coastal dry and wet savannas appears to be vulnerable to plant invasion (with 165 naturalized species, about 34% of the alien flora), especially by Acacia mangium (Mimosaceae) and Melaleuca quinquenervia (Myrtaceae) which are forming localized but dense monotypic stands. Both tree species, intentionnally introduced for reforestation, rehabilitation, and as garden ornamentals and have the potential to spread with increasing human disturbances The number and abundance of naturalized alien plants in the relatively undisturbed tropical lowland rainforests and savannas remains still very low. Therefore, surveillance, early detection, and eradication of potential plant invaders are crucial; moreover collaboration with neighbouring countries of the Guiana Shield is essential to prevent the introduction of potentially invasive species which are still not present in French Guiana.     

Naturalization of introduced plants: ecological drivers of biogeographical patterns

CONTENTS: Summary 383 I. Introduction 383 II. The introduction-naturalization-invasion continuum for conceptualizing biological invasions 384 III. The biogeographical background for studying naturalization: variation among populations and regions 385 IV. Factors determining naturalization in plants 388 Acknowledgements 392 References 392 SUMMARY: The literature on biological invasions is biased in favour of invasive species - those that spread and often reach high abundance following introduction by humans. It is, however, also important to understand previous stages in the introduction-naturalization-invasion continuum ('the continuum'), especially the factors that mediate naturalization. The emphasis on invasiveness is partly because most invasions are only recognized once species occupy large adventive ranges or start to spread. Also, many studies lump all alien species, and fail to separate introduced, naturalized and invasive populations and species. These biases impede our ability to elucidate the full suite of drivers of invasion and to predict invasion dynamics, because different factors mediate progression along different sections of the continuum. A better understanding of the determinants of naturalization is important because all naturalized species are potential invaders. Processes leading to naturalization act differently in different regions and global biogeographical patterns of plant invasions result from the interaction of population-biological, macroecological and human-induced factors. We explore what is known about how determinants of naturalization in plants interact at various scales, and how their importance varies along the continuum. Research that is explicitly linked to particular stages of the continuum can generate new information that is appropriate for improving the management of biological invasions if, for example, potentially invasive species are identified before they exert an impact.     

Macroecological drivers of alien conifer naturalizations worldwide

Understanding the factors that drive the global distribution of alien species is a pivotal issue in invasion biology. Here, we used data on naturalized conifers (Pinaceae, Cupressaceae) from sixty temperate and subtropical regions and five continents to test how environmental and socio‐economic conditions of recipient areas as well as introduction efforts affect naturalization probabilities. We collated 18 predictor variables for each region describing environmental, biogeographic and socio‐economic conditions as well as a measure of the macro‐climatic match with the species' native ranges, and the extent to which alien conifers are used in commercial forestry. Naturalization probabilities across all species and regions were then related to these predictor variables by means of generalized linear mixed models. For both Pinaceae and Cupressaceae, naturalization probabilities were generally higher in the Southern Hemisphere, and increased with indicators of habitat diversity of the recipient region. The match in macro‐climatic conditions between the native and introduced regions was a significant predictor of conifer naturalization, but socio‐economic variables were less powerful predictors. Only for Cupressaceae did a socio‐economic variable (human population density) affect naturalization probabilities. Key attributes facilitating naturalization were related to introduction effort. Moreover, usage in commercial forestry generally fostered naturalization, although the actual size of alien conifer plantations in a region was only correlated with the naturalization of Pinaceae. Our results suggest that climate matching, habitat diversity and introduction effort co‐determine the probability of naturalization, which additionally, is modulated by biogeographic features of the recipient area, such as incidence of natural enemies or competitors. To date, the most widely used tools for invasive plant risk assessment only account for climate match and rarely factor in other attributes of the recipient environment. Future tools should additionally consider biotic environment and introduction effort if risk assessment is to be effective.     

Predicting the spread of an invasive plant: combining experiments and ecological niche model

Rapid evolutionary adjustments to novel environments may contribute to the successful spread of invasive species, and can lead to niche shifts making range dynamics unpredictable. These effects might be intensified by artificial selection in the course of breeding efforts, since many successful plant invaders were deliberately introduced and cultivated as ornamentals. We hypothesized that the invasion success of Buddleja davidii, the ornamental butterfly bush, is facilitated by local adaptation to minimum temperatures and thus, exhibits unpredictable range dynamics. To assess the potential effects of adaptive evolution and artificial selection on the spread of B. davidii, we combined a common garden experiment investigating local adaptation to frost, with ecological niche modelling of the species’ native and invasive ranges. We expected that populations naturalized in sub‐continental climate are less susceptible to frost than populations from oceanic climate, and that the invasive range does not match predictions based on climatic data from the native range. Indeed, we revealed significant variation among invasive B. davidii populations in frost resistance. However, frost hardiness was not related to geographic location or climatic variables of the populations’ home site, suggesting that invasive B. davidii populations are not locally adapted to minimum temperatures. This is in line with results of our ecological niche model that did not detect a niche shift between the species’ native range in China, and its invasive range in Europe and North America. Furthermore, our niche model showed that the potential invasive range of B. davidii is still not completely occupied. Together with the frost resistance data obtained in our experiment, the results indicate that climatic conditions are currently not limiting the further spread of the species in Europe and North America.     

Loci under selection during multiple range expansions of an invasive plant are mostly population specific,but patterns are associated with climate

Identifying the genes underlying rapid evolutionary changes, describing their function and ascertaining the environmental pressures that determine fitness are the central elements needed for understanding of evolutionary processes and phenotypic changes that improve the fitness of populations. It has been hypothesized that rapid adaptive changes in new environments may contribute to the rapid spread and success of invasive plants and animals. As yet, studies of adaptation during invasion are scarce, as is knowledge of the genes underlying adaptation, especially in multiple replicated invasions. Here, we quantified how genotype frequencies change during invasions, resulting in rapid evolution of naturalized populations. We used six fully replicated common garden experiments in Brazil where Pinus taeda (loblolly pine) was introduced at the same time, in the same numbers, from the same seed sources, and has formed naturalized populations expanding outward from the plantations. We used a combination of nonparametric, population genetics and multivariate statistics to detect changes in genotype frequencies along each of the six naturalization gradients and their association with climate as well as shifts in allele frequencies compared to the source populations. Results show 25 genes with significant shifts in genotype frequencies. Six genes had shifts in more than one population. Climate explained 25% of the variation in the groups of genes under selection across all locations, but specific genes under strong selection during invasions did not show climate‐related convergence. In conclusion, we detected rapid evolutionary changes during invasive range expansions, but the particular gene‐level patterns of evolution may be population specific.     

Is the Climate Right for Pleistocene Rewilding? Using Species Distribution Models to Extrapolate Climatic Suitability for Mammals across Continents

Species distribution models (SDMs) are increasingly used for extrapolation, or predicting suitable regions for species under new geographic or temporal scenarios. However, SDM predictions may be prone to errors if species are not at equilibrium with climatic conditions in the current range and if training samples are not representative. Here the controversial “Pleistocene rewilding” proposal was used as a novel example to address some of the challenges of extrapolating modeled species-climate relationships outside of current ranges. Climatic suitability for three proposed proxy species (Asian elephant, African cheetah and African lion) was extrapolated to the American southwest and Great Plains using Maxent, a machine-learning species distribution model. Similar models were fit for Oryx gazella, a species native to Africa that has naturalized in North America, to test model predictions. To overcome biases introduced by contracted modern ranges and limited occurrence data, random pseudo-presence points generated from modern and historical ranges were used for model training. For all species except the oryx, models of climatic suitability fit to training data from historical ranges produced larger areas of predicted suitability in North America than models fit to training data from modern ranges. Four naturalized oryx populations in the American southwest were correctly predicted with a generous model threshold, but none of these locations were predicted with a more stringent threshold. In general, the northern Great Plains had low climatic suitability for all focal species and scenarios considered, while portions of the southern Great Plains and American southwest had low to intermediate suitability for some species in some scenarios. The results suggest that the use of historical, in addition to modern, range information and randomly sampled pseudo-presence points may improve model accuracy. This has implications for modeling range shifts of organisms in response to climate change.     

The biogeography of naturalization in alien plants

Aim  This paper reviews the main geographical determinants of naturalization in plants.
Location  Global.
Methods  Comparative studies of large data sets of alien floras are the main source of information on global patterns of naturalization.
Results  Temperate mainland regions are more invaded than tropical mainland regions but there seems to be no difference in invasibility of temperate and tropical islands. Islands are more invaded than the mainland. The number of naturalized species in temperate regions decreases with latitude and their geographical ranges increase with latitude. The number of naturalized species on islands increases with temperature. Naturalized species contribute to floristic homogenization, but the phenomenon is scale-dependent.
Main conclusions  Some robust patterns are evident from currently available data, but further research is needed on several aspects to advance our understanding of the biogeography of naturalization of alien plants. For example, measures of propagule pressure are needed to determine the invasibility of communities/ecosystems/regions. The patterns discussed in this paper are derived largely from numbers and proportions of naturalized species, and little is known about the proportion of introduced species that become naturalized. Further insights on naturalization rates, i.e. the proportion of aliens that successfully naturalize within regions, and on geographical and other determinants of its variation would provide us with better understanding of the invasion process. Comparative studies, and resulting generalizations, are almost exclusively based on numbers of species, but alien species differ in their impact on native biodiversity and ecosystem processes.     

区域种植业气候适宜度及其对种植活动的响应——以四川省盐亭县为例

在分析盐亭县近63年来(1950—2012)种植业生产发展的基础上,选取该县农村社会经济条件相对稳定的近32年(1981—2012)为研究时段。运用农业生态气候适宜度方法,依据水稻、红薯、玉米、小麦和油菜等5种主要作物生育期的光、热、水等气候条件,分别估算各种作物的资源适宜指数、效能适宜指数和利用指数,构建小尺度区域种植业气候适宜度模型和种植活动对区域种植业气候适宜度的影响度模型,进行小尺度区域种植业气候适宜度以及种植活动对种植业气候适宜度的影响度估算,并对种植业生产对气候变化的适应进行探讨。研究结果表明,(1)近32年来盐亭县大春作物的平均资源适宜指数、效能适宜指数和利用指数(分别为0.578、0.281和48.37%)均大于小春作物(分别为0.304、0.128和42.24%),大春作物的气候适宜度高于小春作物,且作物间的气候适宜度差异较大。(2)受季风气候波动的影响,该县作物气候适宜度有明显的年际波动;该县近32年来气候变化对大春作物气候适宜度有轻微不利影响,而对小春作物气候适宜度趋于有利。(3)盐亭县近32年来种植业平均的资源适宜指数为0.466、效能适宜指数为0.212、利用指数为45.49%;受5种作物资源适宜指数、效能适宜指数,以及作物播种面积与产量年际波动的综合影响,该县种植业气候适宜度亦有明显的年际波动;气候变化对该县种植业气候适宜度总体上有不利影响。(4)近32年来该县种植活动对种植业气候适宜度的影响度平均值为0.00092,其年际波动较大。通过作物种植组合结构的调整,在20世纪90年代中期前对种植业气候适宜度的提高有微弱的正向影响,对气候变化有一定程度的适应;而后期则有负向作用。     

Cultivation Fosters Plant Naturalization by Reducing Environmental Stochasticity

A large fraction of the immigrant (or founder) populations of terrestrial plants are small (< 10⁴) and are acutely sensitive to environmental stochasticity. As a result, they undergo radical size fluctuations during a prolonged lag phase that almost always result in their extirpation. Naturalizations are those rare examples in which an immigrant population increases above a threshold size such that the consequences of environmental stochasticity are markedly lower. The likelihood that a non-indigenous population will reach this threshold size would be enhanced substantially through either deliberate or inadvertent cultivation. Cultivation (e.g. protection from predators, parasites, drought, frost) shields small immigrant populations from the extreme expressions of environmental stochasticity. In addition, cultivation can preserve through seed storage a residual non-indigenous population from which new populations can be established. As disseminules are spread locally, and even regionally, immigrant populations sample a wide variety of micro-habitats, thus increasing the likelihood that some plants will survive even without cultivation. Origins of naturalized floras in Australia and the US reveal a strong circumstantial link between cultivation and subsequent naturalization: the single largest group of naturalized species was deliberately introduced as either crops, forage spp., or ornamentals. Another group was introduced inadvertently as contaminants in crop seeds. This correspondence between cultivation and subsequent naturalization provides a common demographic explanation for non-indigenous plant persistence that largely transcends species’ attributes and the commonly ascribed features of communities that are vulnerable to the entry of non-indigenous plants. Humans have played a more profound role in fostering plant naturalizations than by acting simply as plant dispersers; their post-immigration cultivation fosters much naturalization. This revised version was published online in July 2006 with corrections to the Cover Date.     

Patterns of plant invasions in China: Taxonomic,biogeographic, climatic approaches and anthropogenic effects

This study was aimed to determine the patterns as well as the effects of biological, anthropogenic, and climatic factors on plant invasions in China. About 270 volumes of national and regional floras were employed to compile a naturalized flora of China. Habit, life form, origin, distribution, and uses of naturalized plants were also analyzed to determine patterns on invasion. Correlations between biological, anthropogenic and climatic parameters were estimated at province and regional scales. Naturalized species represent 1% of the flora of China. Asteraceae, Fabaceae, and Poaceae are the dominant families, but Euphorbiaceae and Cactaceae have the largest ratios of naturalized species to their global numbers. Oenothera, Euphorbia, and Crotalaria were the dominant genera. Around 50% of exotic species were introduced intentionally for medicinal purposes. Most of the naturalized species originated in tropical America, followed by Asia and Europe. Number of naturalized species was significantly correlated to the number of native species/log area. The intensity of plant invasion showed a pattern along climate zones from mesic to xeric, declining with decreasing temperature and precipitation across the nation. Anthropogenic factor, such as distance of transportation, was significantly correlated to plant invasions at a regional scale. Although anthropogenic factors were largely responsible for creating opportunities for exotic species to spread and establish, the local biodiversity and climate factors were the major factors shaping the pattern of plant invasions in China. The warm regions, which are the hot spots of local biodiversity, and relatively developed areas of China, furthermore, require immediate attentions.     

How strongly do interactions with closely‐related native species influence plant invasions? Darwin's naturalization hypothesis assessed on Mediterranean islands

Aim Recent works have found the presence of native congeners to have a small effect on the naturalization rates of introduced plants, some suggesting a negative interaction (as proposed by Charles Darwin in The Origin of Species), and others a positive association. We assessed this question for a new biogeographic region, and discuss some of the problems associated with data base analyses of this type. Location Islands of the Mediterranean basin. Methods Presence or absence of congeners was assessed for all naturalized alien plants species at regional, local and habitat scales. Using general linear models, we attempted to explain the abundance of the species (as measured by the number of islands where recorded) from their congeneric status, and assessed whether the patterns could be alternatively accounted for by a range of biological, geographical and anthropogenic factors. A simulation model was also used to investigate the impact of a simple bias on a comparable but hypothetical data set. Results Data base analyses addressing Darwin's hypothesis are prone to bias from a number of sources. Interaction between invaders and congenerics may be overestimated, as they often do not co‐occur in the same habitats. Furthermore, intercorrelations between naturalization success and associated factors such as introduction frequency, which are also not independent from relatedness with the native flora, may generate an apparent influence of congenerics without implying a biological interaction. We detected no true influence from related natives on the successful establishment of alien species of the Mediterranean. Rarely‐introduced species tended to fare better in the presence of congeners, but it appears that this effect was generated because species introduced accidentally into highly invasible agricultural and ruderal habitats have many relatives in the region, due to common evolutionary origins. Main conclusions Relatedness to the native flora has no more than a marginal influence on the invasion success of alien plants in the Mediterranean, although apparent trends can easily be generated through artefacts of the data base.     

中国归化植物组成特征及其时空分布格局分析

入侵植物是归化植物的子集, 归化是植物入侵的基本前提, 完整的归化植物清单有助于预测和预防植物入侵。该文基于实地调查和资料收集, 对中国归化植物种类及分布进行统计, 结果表明, 当前中国共有归化植物112科578属1 099种, 约70%为人为引入, 而美洲是其最大原产地, 植物生活型以草本为主, 分类学组成中菊科、禾本科、豆科最多。在空间尺度上, 物种多样性及密度均呈自东南沿海向西北内陆递减的趋势, 纬度和国内生产总值是其空间分布格局的主要驱动因素, 被殖民史、植物园分布及调查次数在一定程度上影响空间分布格局; 在时间尺度上, 归化植物物种多样性当前正处在快速增长阶段并可能持续20-30年, 社会经济发展、人为干扰、土地利用方式改变及全球变化是其主要驱动因素。该研究丰富了中国归化植物的本底资料, 有助于了解外来植物入侵的形式与威胁, 并为外来植物综合管控或本地生物多样性保护提供依据。     

Naturalization of exotic plant species in north‐eastern North America: trends and detection capacity

Aim To reconstruct historical naturalization trends of exotic vascular plants in Quebec and Massachusetts, two regions that experienced fluctuations in their herbarium specimen collection efforts. We hypothesized that the observed trends are linked to the collection effort and that they differ according to the motive for plant introduction. Methods A checklist of naturalized exotic plant taxa for Quebec, including oldest proof of naturalization and motive for introduction, was built from herbaria, journals, dissertations, theses, bulletins, websites and unpublished records. For Massachusetts, we used the checklist of Sorrie (2005; Rhodora, 107 , 284–329). The collection effort was estimated from a pool of species with specimens from the five largest Quebec herbaria and from the Harvard University Herbarium. Results Naturalization trends of exotic plants in Quebec are similar to those observed in Massachusetts. A large part of the plants naturalized in the 17th, 18th or 19th centuries, with very little naturalization during the last 30–90 years. In general, there is a significant statistical link between herbarium specimen collection effort and the discovery of new naturalized plant taxa. Very few utilitarian plants naturalized in either region during the 20th century. In Quebec, a high number of ornamental plants naturalized during the last 100 years. Main conclusions The link between herbarium specimen collection effort and the discovery of new naturalized plants is real, although not as straightforward as it seems. Our analysis suggests that at least part of the decline in the number of new naturalized exotic plants observed in Quebec and Massachusetts during the last 30–90 years is a direct consequence of the low interest for traditional floristic studies. However, the possibility of a real decline cannot be ruled out. We nevertheless provide here one of the first pieces of evidence of the potential consequences of the decline of local plant collection for environmental management and especially for early detection systems of new invaders.     

Estimating the invasion success of introduced plants

We present methods for estimating the base proportion of introduced alien species that will naturalize, and the distribution of time until naturalization occurs. The approach is Bayesian, incorporating prior estimates of the probability of naturalization and the time from introduction to naturalization. A worked example uses data on the introduction and time to recorded naturalization of woody perennials introduced to South Australia. Up until 2007, 188 of 2230 (8.4%) woody perennials listed in nursery catalogues between 1843 and 1985 were recorded as having naturalized. If prior information on naturalization rates from elsewhere is ignored, the available data suggest that the most likely proportion of introduced plants that will naturalize is large (0.93) though uncertain (95% CI 0.51–0.99), with the corresponding mean time to recorded naturalization being protracted (522 years) and similarly uncertain (95% CI 360–678 years). Alternatively, if informative prior estimates of both the naturalization probability and the time to recorded naturalization are incorporated, the most likely probability of naturalization is estimated to be 18.6% (95% CI 15.5–23.4%). For those plants that do naturalize, the most likely value for the mean time from importation to recorded naturalization is 149 years (95% CI 130–174 years). Our results illustrate the potentially long timescale of the naturalization process, and the challenges this presents for obtaining accurate estimates of naturalization parameters.     

Naturalization of alien plants in China

Naturalization (the establishment of a self-sustaining population for at least a decade) is a fundamental precondition for plant invasion and so compiling a complete inventory of naturalized alien species is necessary for predicting and hence preventing such invasion. However, nationwide information on naturalized plants in China is still lacking. We compiled a nationwide list of the naturalized plant species of China, based on various literature reports. The list comprised a total of 861 naturalized plant species belonging to 110 families and 465 genera. The three most dominant families were Compositae, Poaceae, and Leguminosae, accounting for 16, 13 and 12% of naturalized plants, respectively. Among genera, Euphorbia and Solanum had the most naturalized species, followed by Ipomoea, Amaranthus, Oenothera, and Trifolium. Over half of all aliens were of American origin (52%), followed by those with European (14%) and Asian (13%) origins. Annuals and perennial herbs were prevalent among naturalized species; comparison to other studies suggests however that the invasive potential is higher among plants with longer life cycles than those of annuals. The taxonomic pattern of plant naturalization in China is similar to patterns worldwide. However, the low proportion of naturalized plants within the Chinese flora overall suggests that the potential for plant invasions in China may be high. Therefore, greater attention should be focused on naturalization of alien plants in China, especially concerning species of dominant families or genera, and those with a perennial life cycle.     

Diploid male production in a rare and locally distributed bumblebee, Bombus florilegus (Hymenoptera, Apidae)

The European bumblebee B. terrestris was recently introduced in Japan for agricultural purposes and has now become naturalized. The naturalization of this exotic species may have great detrimental effects on closely related native Japanese bumblebees. The Japanese bumblebee Bombus florilegus is a rare and locally distributed species found in the Nemuro Peninsula of Hokkaido, Japan. In order to assess its population genetics, we estimated the genetic structure of B. florilegus in 16 breeding colonies (queen, workers, and males) and 20 foraging queens by analyzing microsatellite DNA markers. Of the 36 queens analyzed by genotyping and dissection, 32 had been inseminated by a male. The remaining 4 had not been inseminated at all. Of the 4 nonmated queens, one was triploid. Diploid males were found in 4 breeding colonies. Based on the microsatellite data, it appears that B. florilegus has low reproductive success. Since matched mating and nonmating within local populations are high, the extinction risk is correspondingly higher. Our results suggest that conservation of the Japanese B. florilegus is required in order to protect it from both habitat destruction and the naturalization of alien species. Received 19 July 2007; revised 13 October 2007; accepted 15 October 2007.     

The role of adaptive strategies in plant naturalization

Determining the factors associated with the naturalization of alien species is a central theme in ecology. Here, we tested the usefulness of a metric for quantifying Grime's seminal concept of adaptive strategies – competitors, stress‐tolerators and ruderals (CSR) – to explain plant naturalizations worldwide. Using a global dataset of 3004 vascular plant species, and accounting for phylogenetic relatedness and species’ native biomes, we assessed the associations between calculated C‐, S‐ and R‐scores and naturalization success for species exhibiting different life forms. Across different plant life forms, C‐scores were positively and S‐scores negatively associated with both the probability of naturalization and the number of regions where the species has naturalized. R‐scores had positive effects on the probability of naturalization. These effects of the scores were, however, weak to absent for tree species. Our findings demonstrate the utility of CSR‐score calculation to broadly represent, and potentially explain, the naturalization success of plant species.