The naturalization to invasion transition: Are there introduction‐history correlates of invasiveness in exotic plants of Australia?

Of the large number of exotic plant species that become naturalized in new geographic regions, only a subset make the transition to become invasive. Identifying the factors that underpin the transition from naturalization to invasion is important for our understanding of biological invasions. To determine introduction‐history correlates of invasiveness among naturalized plant species of Australia, we compared geographic origin, reason for introduction, minimum residence time and growth form between naturalized non‐invasive species and naturalized invasive plant species. We found that more invasive species than expected originated from South America and North America, while fewer invasive species than expected originated from Europe and Australasia. There was no significant difference between invasive and non‐invasive species with respect to reason for introduction to Australia. However, invasive species were significantly more likely to have been resident in Australia for a longer period of time than non‐invasive species. Residence times of invasive species were consistently and significantly higher than residence times of non‐invasive species even when each continent of origin was considered separately. Furthermore, residence times for both invasive and non‐invasive species varied significantly as a function of continent of origin, with species from South America having been introduced to Australia more recently on average than species from Europe, Australasia and North America. We also found that fewer invasive species than expected were herbs and more invasive species than expected were primarily climbers. Considered together, our results indicate a high propensity for invasiveness in Australia among exotic plant species from South America, given that they appear in general capable of more rapid shifts to invasiveness than aliens from other regions. Furthermore, our findings support an emerging global generality that introduction‐history traits must be statistically controlled for in comparative studies exploring life‐history and ecological correlates of invasion success.     

Invasiveness and homogenization: synergism of wide dispersal and high local abundance

Aim To determine whether invasive and locally abundant non‐native species have a more homogenizing effect on plant communities than non‐invasive and less abundant non‐native species. Location California and Florida counties, conservation areas in the USA, and eight US cities. Methods Species lists among counties, conservation areas and cities were compared to see whether invasive and abundant non‐native species increased the Jaccard index of similarity between localities beyond any increases caused by non‐invasive and less abundant non‐native species. Results For all comparisons, we found that invasive non‐native species have a significantly greater homogenizing effect than non‐invasive non‐native species. For the US conservation areas, we found that locally abundant invasive species tend to be more widespread and more widely shared than less abundant invasive species. There is also a positive relationship between homogenization by invasive species and the magnitude of human disturbance. Main conclusions Invasive non‐native species tend to be disproportionately shared among communities relative to non‐invasive non‐native species. This effect is enhanced by human disturbance, as measured by the ratio of non‐native to native species. There is a synergism between abundance and geographical range which enhances the homogenizing effects of abundant species. Invasive species, with wide ecological niches, are more widely shared among communities and more locally abundant. Abundant invasive species are thus more spatially homogenizing, and more ecologically dominant (functionally homogenizing). Also, ‘perceived homogenization’ is probably greater than homogenization measured by the increase in shared species. The abundant species typically seen by the casual observer in a biological community are probably more commonly shared between communities than less common species. Studies that lack abundance data and measure homogenization only on the basis of shared species, which includes most homogenization studies to date, probably underestimate the homogenizing impacts of non‐native species as perceived by people.     

Preadapted for invasiveness: do species traits or their plastic response to shading differ between invasive and non‐invasive plant species in their native range?

Aim Species capable of vigorous growth under a wide range of environmental conditions should have a higher chance of becoming invasive after introduction into new regions. High performance across environments can be achieved either by constitutively expressed traits that allow for high resource uptake under different environmental conditions or by adaptive plasticity of traits. Here we test whether invasive and non‐invasive species differ in presumably adaptive plasticity. Location Europe (for native species); the rest of the world and North America in particular (for alien species). Methods We selected 14 congeneric pairs of European herbaceous species that have all been introduced elsewhere. One species of each pair is highly invasive elsewhere in the world, particularly so in North America, whereas the other species has not become invasive or has spread only to a limited degree. We grew native plant material of the 28 species under shaded and non‐shaded conditions in a common garden experiment, and measured biomass production and morphological traits that are frequently related to shade tolerance and avoidance. Results Invasive species had higher shoot–root ratios, tended to have longer leaf‐blades, and produced more biomass than congeneric non‐invasive species both under shaded and non‐shaded conditions. Plants responded to shading by increasing shoot–root ratios and specific leaf area. Surprisingly, these shade‐induced responses, which are widely considered to be adaptive, did not differ between invasive and non‐invasive species. Main conclusions We conclude that high biomass production across different light environments pre‐adapts species to become invasive, and that this is not mediated by plasticities of the morphological traits that we measured.     

Are Hong Kong and Taiwan stepping‐stones for invasive species to the mainland of China?

Understanding the origins and introduction pathways of invasive species is a fundamental issue for invasion biology, which is necessary for predicting and preventing future invasion. Once an invasive species is established in a new location, this location could serve as a stepping‐stone for further invasions. However, such “stepping‐stone” effect has not been widely investigated. Using the published literature and records, we compiled the first found locations of 127 top invasive species in China. Our study showed that the most common landing spots of these invasive species were Hong Kong (22 species) and Taiwan (20 species), which accounted for one‐third of the invasive species in China. Our analysis revealed that the invasive species in mainland China were more likely to transport from Hong Kong than Macau, a neighboring region with a similar area and colonial history. Similarly, more invasive species were also first landed on Taiwan than Hainan, a nearby island sharing similar climate conditions. Together, our findings indicate that Hong Kong and Taiwan are the most important stepping‐stones for invasive species to the mainland of China and suggesting that the increasing trade exchange of China's coastal ports constitutes a potential risk for the spread of more invasive species. We suppose that they would be the future stepping‐stones for invasive species to the mainland of China and these coastal ports regions where improved biosecurity is needed now.     

Little evidence for release from herbivores as a driver of plant invasiveness from a multi‐species herbivore‐removal experiment

Enemy release is frequently posed as a main driver of invasiveness of alien species. However, an experimental multi‐species test examining performance and herbivory of invasive alien, non‐invasive alien and native plant species in the presence and absence of natural enemies is lacking. In a common garden experiment in Switzerland, we manipulated exposure of seven alien invasive, eight alien non‐invasive and fourteen native species from six taxonomic groups to natural enemies (invertebrate herbivores), by applying a pesticide treatment under two different nutrient levels. We assessed biomass production, herbivore damage and the major herbivore taxa on plants. Across all species, plants gained significantly greater biomass under pesticide treatment. However, invasive, non‐invasive and native species did not differ in their biomass response to pesticide treatment at either nutrient level. The proportion of leaves damaged on invasive species was significantly lower compared to native species, but not when compared to non‐invasive species. However, the difference was lost when plant size was accounted for. There were no differences between invasive, non‐invasive and native species in herbivore abundance. Our study offers little support for invertebrate herbivore release as a driver of plant invasiveness, but suggests that future enemy release studies should account for differences in plant size among species.     

Invasion success and threat status: two sides of a different coin?

The characteristics possessed by invasive species have been suggested to be the reverse of those possessed by species threatened with extinction, such that relationships of species’ traits to invasion success should be opposite in sign to relationships of the same traits to extinction threat. A recent study (Jeschke, J. M. and Strayer, D. L. 2008. Are threat status and invasion success two sides of the same coin? – Ecography 31: 124–130) found no evidence for this “two‐sides‐of‐the‐same‐coin” hypothesis but compared characteristics of species in each taxon that were invasive to a control group consisting of all other species. A different view of the “two‐sides‐of‐the‐same‐coin” hypothesis may be obtained if the characters of invasive species are compared to those of a control group consisting of species that have not invaded despite actually being introduced. Here, we show that changing the control group for comparison with invasive species does not change the lack of support for the “two‐sides‐of‐the‐same‐coin” hypothesis but does change views about which specific traits are consistent with the hypothesis.     

Non‐additive effects of invasive tree litter shift seasonal N release: a potential invasion feedback

Many invasive plant species strongly alter ecosystem processes by producing leaf litter that decomposes faster and releases N more quickly than that of native species. However, while most studies of invasive species litter impacts have only considered the decomposition of species in monoculture, forest litter layers typically contain litter from many species. Many litter mixtures decompose in a non‐additive manner, in which the mixture decomposes more quickly (synergistic effect) or more slowly (antagonistic effect) than would be expected based on decomposition of the component species’ litters in isolation. We investigated the potential for non‐additive effects of invasive species’ litter by conducting a one‐year litter bag experiment in which we mixed the litters of four native tree species with each of four invasive species. Litter mixtures frequently lost mass at non‐additive rates, although not at every loading ratio, and the presence, sign, and strength of effects depended on species composition. Non‐additive effects on N loss occurred in more litter combinations, and were almost always antagonistic at 90 days and synergistic at 365 days. Invasive species litter with lower C:N led to more strongly synergistic N loss with time. During the growing season, non‐additive patterns of N loss almost always resulted in increased N release – up to six times greater than would be expected based on single‐species decomposition. Consequently, we suggest that invasive species may further synchronize N release from the litter layer with plant N demand, enhancing any positive litter feedback to invasion. These results highlight the need to consider non‐additive effects of litter mixing in invaded forest communities, and suggest that estimates of invasive species’ impacts on ecosystem processes would be improved by considering these effects.     

Profiling invasive fish species: the importance of phylogeny and human use

Understanding the ecological differences between native and invasive species is of considerable scientific and practical interest. We examined such differences between native and invasive inland fish species from the Iberian Peninsula in order to analyse the importance of phylogenetic correction and variability (in addition to central tendency). We collected 26 quantitative and qualitative variables on the ecology, life‐history traits and human use of the 69 inland fish species of the Iberian Peninsula, including native, invasive and migratory species. The taxonomic distribution of invasive fish species deviated significantly from world freshwater richness and in contrast to native species, invasive fish belongs to only five taxonomic orders but to a wide spectrum of families not native to the Iberian Peninsula. Because the life‐history traits were highly dependent on taxonomy, the results, with or without applying phylogenetic methods, differed and after accounting for phylogeny, invasive species displayed higher and wider latitude in general and a different reproductive season mainly among salmonids and cyprinids. Human use was also significantly different between native and invasive fish species and produced more variability in life‐history traits of invasive species and uneven taxonomic distribution because of the high diversity of species introduced. We show that accounting for taxonomy and studying variability in addition to central tendency is important in the comparison of life‐history traits between native and invasive species.     

Revisiting the use of the invasive species concept: An empirical approach

Invasion science has not been developed without controversies. Two questions that are still unsolved are: what is an invasive species?, and are invasive species an inherent conservation problem? These questions have led to discussions about effects versus origins. In contrast to the definitional problems, a unified framework describing invasion as a step‐by‐step process has been widely accepted. I conducted a bibliographic search with two separate databases searching for (i) evidence of less use of controversial terms over time; (ii) how many articles defined ‘invasive species’; (iii) the criteria used to define a species as invasive; and (iv) in which stage of the invasion continuum were species labelled as invasive located. My results show that controversial terms are widely used, that authors rarely define ‘invasive species’ and, often, it is very complicated to determine which criterion they used. In addition, only a fraction of the species labelled as invasive could be classified as such according to the unified framework of invasion stages. This is not a merely semantic issue, because invasive is a strong and value‐laden term that is used to guide environmental agendas. The uncritical use of a key concept could hamper research, complicate communication among peers and produce mixed results.     

Do invasive freshwater fish species grow better when they are invasive?

A frequent assumption in invasion ecology is that invasive species have enhanced growth rates in their invasive ranges. However, invasions frequently occur in sub‐tropical and tropical environments where growth could be higher simply due to climatic conditions rather than novel habitat. In this study, a meta‐analysis of growth rates (length‐at‐age data) was completed for six invasive freshwater fish species: common carp Cyprinus carpio, largemouth bass Micropterus salmoides, brown trout Salmo trutta, brown bullhead Ictalurus nebulosus, flathead catfish Pylodictis olivaris and northern snakehead Channa argus. Significant effects of climate on growth were observed for all species except common carp, and following normalization of growth for climate effects, a range of growth responses between native and invasive populations were revealed. Two species (brown trout, flathead catfish) showed significantly increased growth rates in invasive compared to native ranges, but two species (common carp, largemouth bass) showed significantly faster growth in native ranges, and two other species (northern snakehead, brown bullhead) showed no difference in growth rates. No species showed both significantly enhanced growth rates and initial sizes in invasive compared to native ranges. Using the comparative method, countergradient growth variations were apparent for all species within their native ranges and for all but one species in invasive ranges. Invasive populations of freshwater fish do not always grow faster when invasive and future studies need to consider growth covariates (e.g. climate and countergradient growth) prior to comparing life‐history differences between invasive and native populations.     

Reproductive output of invasive versus native plants

Aim Propagule size and output are critical for the ability of a plant species to colonize new environments. If invasive species have a greater reproductive output than native species (via more and/or larger seeds), then they will have a greater dispersal and establishment ability. Previous comparisons within plant genera, families or environments have conflicted over the differences in reproductive traits between native and invasive species. We went beyond a genus‐, family‐ or habitat‐specific approach and analysed data for plant reproductive traits from the global literature, to investigate whether: (1) seed mass and production differ between the original and introduced ranges of invasive species; (2) seed mass and production differ between invasives and natives; and (3) invasives produce more seeds per unit seed mass than natives. Location Global. Methods We combined an existing data set of native plant reproductive data with a new data compilation for invasive species. We used t‐tests to compare original and introduced range populations, two‐way ANOVAs to compare natives and invasives, and an ANCOVA to examine the relationship between seed mass and production for natives and invasives. The ANCOVA was performed again incorporating phylogenetically independent contrasts to overcome any phylogenetic bias in the data sets. Results Neither seed mass nor seed production of invasive species differed between their introduced and original ranges. We found no significant difference in seed mass between invasives and natives after growth form had been accounted for. Seed production was greater for invasive species overall and within herb and woody growth forms. For a given seed mass, invasive species produced 6.7‐fold (all species), 6.9‐fold (herbs only) and 26.1‐fold (woody species only) more seeds per individual per year than native species. The phylogenetic ANCOVA verified that this trend did not appear to be influenced by phylogenetic bias within either data set. Main conclusions This study provides the first global examination of both seed mass and production traits in native and invasive species. Invasive species express a strategy of greater seed production both overall and per unit seed mass compared with natives. The consequent increased likelihood of establishment from long‐distance seed dispersal may significantly contribute to the invasiveness of many exotic species.     

A risk‐based inventory of invasive plant species of Queensland,Australia: Regional,ecological and floristic insights

Invasive alien plant species threaten agriculture and biodiversity globally and require ongoing management to minimise impacts. However, the large number of invasive species means that a risk‐based approach to prioritisation is needed, taking into account the spatial scale of management decisions and myriad of available information. Here, we developed a risk‐based inventory of invasive plants in Queensland, Australia, using both current species distribution/abundance and the severity of their impacts. Our assessment followed a comprehensive data collection process including a scoping of local government pest management plans, herbarium records, the published literature and structured elicitation of expert knowledge during a series of regional stakeholder workshops. From ~300 plant species that were identified as established and/or emerging invaders in the State, only one‐third were considered by practitioners to pose significant risks across regions to be considered management priorities. We aggregated regional species lists into a statewide priority list and analysed the data set (107 species) for historical, geographical, floristic and ecological patterns. Regions on the mainland eastern seaboard of the State share similar invasive plant communities, suggesting that these regions may form a single management unit, unlike the western/inland and the extreme far north (Torres Strait Islands) regions, which share fewer invasive plant species. Positive correlations were detected between invasiveness and time since introduction for some but not all plant life forms. Stakeholders identified research and management priorities for the invasive plant list, including biological control options, public awareness/education, effective herbicide use, ecology/taxonomy and risk analysis. In the course of the exercise, a statewide invasive plant priority list of high‐, medium‐ and low‐impact scores for policy, research and management was compiled. Finally, our approach to invasive plant species prioritisation highlighted that planning and policy documents are not necessarily reflected at the grass‐root level in terms of species identity and management priorities.     

Invasive alien plant species in China: regional distribution patterns

Plant invasions have been attracting increasing attention from ecologists because of their worldwide environmental impacts and huge economic costs. Research on the characteristics of the recipient regions is essential for understanding the process of plant invasion. However, few previous studies on invasibility of habitats include social factors, although human activities are critical in the process of plant invasion. China is a vast country with high plant species diversity and a long history of introduction of exotic plant species and is particularly vulnerable to invasive plant species. Alien plant species are widespread in the country. Therefore, the study of invasive plants in China is urgent in practice and theoretically important for developing invasion ecology. For the present study, 126 species were selected to represent the major invasive plant species in China. We then collected data on their species richness in 31 provincial administrative units of China and performed Spearman rank correlations between species richness and possible natural and socio‐economic factors. We found that socio‐economic factors, such as human density and GDP, correlated positively with the species richness of invasive plants in China. In conjunction with the natural and socio‐economic correlations in the study of regional distribution pattern of the major invasive plants, we discussed the factors influencing the regional distribution pattern of the major invasive plants in China. We suggest that native plant species richness was mainly determined by the natural conditions of the regions, while invasive species richness was influenced by natural conditions and human disturbance together.     

A phylogenetic analysis of the British flora sheds light on the evolutionary and ecological factors driving plant invasions

Darwin's naturalization hypothesis predicts that invasive species should perform better in their novel range in the absence of close relatives in the native flora due to reduced competition. Evidence from recent taxonomic and phylogenetic‐based studies, however, is equivocal. We test Darwin's naturalization hypothesis at two different spatial scales using a fossil‐dated molecular phylogenetic tree of the British native and alien flora (ca. 1600 species) and extensive, fine‐scale survey data from the 1998 Countryside Survey. At both landscape and local scales, invasive species were neither significantly more nor less related to the native flora than their non‐invasive alien counterparts. Species invasiveness was instead correlated with higher nitrogen and moisture preference, but not other life history traits such as life‐form and height. We argue that invasive species spread in Britain is hence more likely determined by changes in land use and other anthropogenic factors, rather than evolutionary history. Synthesis. The transition from non‐invasive to invasive is not related to phylogenetic distinctiveness to the native community, but instead to their environmental preferences. Therefore, combating biological invasions in the Britain and other industrialized countries need entirely different strategies than in more natural environments.     

Does localized control of invasive eastern gambusia (Poeciliidae: Gambusia holbrooki) increase population growth of generalist wetland fishes?

While invasive fish management is heavily focussed on containment measures when introductions occur, examples from invasive species management in terrestrial systems suggest that there may also be considerable conservation benefits in implementing localized control programmes. We conducted a field‐based experiment to assess the effectiveness of removing a globally significant invasive fish, eastern gambusia Gambusia holbrooki, from natural wetland habitats of south‐eastern Australia. With recent work suggesting the impacts of eastern gambusia may be minimal for species with generalist life‐history strategies, we hypothesized that the removal of eastern gambusia will reduce localized population growth of the invasive species, but will have little influence on the population growth of more generalist sympatric wetland fish species. We used a predictive modelling approach to investigate changes in eastern gambusia populations following removal activities, and how sympatric fish species responded to such changes. Although eastern gambusia rapidly populated habitats, we demonstrated that control actions substantially reduced the rate of population increase over the four‐month study period. This suggests that control may be an effective localized strategy to suppress eastern gambusia densities. There was however, no evidence of any response to the removal actions by any of the three sympatric fish species investigated – carp gudgeon (Hypseleotris spp.), Australian smelt (Retropinna semoni) and the invasive common carp (Cyprinus carpio). These results support previous work which suggests that the flexible life‐history strategies and behavioural traits of all three species allow co‐existence with eastern gambusia. The study highlights the importance of understanding the potential outcomes of control options which is particularly pertinent for established aquatic invasive species where information on control effectiveness, population dynamics and/or ecosystem response is currently lacking.     

Invasive plants in conservation linkages: a conceptual model that addresses an underappreciated conservation issue

Conservationists have frequently touted the merits of increased landscape connectivity, usually focusing on the efficacy of conservation linkages (corridors) for maintaining viable populations of target species. An often‐mentioned, but still greatly understudied, concern is that increased landscape connectivity via linkages may also aid the movement of undesired species. This paper provides conceptual guidance for research on one major aspect of this gap: invasive plants in conservation linkages. To guide research goals and methods, I develop a conceptual model describing eight interaction types between invasive plants and linkages, i.e. the ways that invasive plants can exist in and move into, through, and out of conservation linkages. Each interaction type within the model has three main components: linkage, matrix, and focal species. I discuss several aspects of these components, including a) differentiating among matrix types, b) understanding edge effects within the linkages, and c) incorporating relevant invasive species’ ecology (primarily dispersal ecology). Spatially‐explicit documentation of invasive plant distribution is essential to understanding these interactions. By focusing on landscape‐scale patterns in real‐world systems, this model will enhance landscape‐level knowledge of invasion ecology and aid land managers in identifying and prioritizing research and management decisions regarding invasive plants in conservation linkages.     

Faster returns on ‘leaf economics’ and different biogeochemical niche in invasive compared with native plant species

Plant‐invasive success is one of the most important current global changes in the biosphere. To understand which factors explain such success, we compared the foliar traits of 41 native and 47 alien‐invasive plant species in Oahu Island (Hawaii), a location with a highly endemic flora that has evolved in isolation and is currently vulnerable to invasions by exotic plant species. Foliar traits, which in most cases presented significant phylogenetic signal, i.e. closely related species tended to resemble each other due to shared ancestry, separated invasive from native species. Invasive species had lower leaf mass per area and enhanced capacities in terms of productivity (photosynthetic capacity) and nutrient capture both of macro‐ (N, P, K) and microelements (Fe, Ni, Cu and Zn). All these differences remain highly significant after removing the effects of phylogenetic history. Alien‐invasive species did not show higher efficiency at using limiting nutrient resources, but they got faster leaf economics returns and occupied a different biogeochemical niche, which helps to explain the success of invasive plants and suggests that potential increases in soil nutrient availability might favor further invasive plant success.     

Pine invasions: climate predicts invasion success; something else predicts failure

Aim Explaining why some invasions fail while others succeed is a prevailing question in invasion biology. Different factors have been proposed to explain the success or failure of exotics. Evidence suggests that climate similarities may be crucial. We tested this using 12 species of the genus Pinus that have been widely planted and shown to be highly invasive. Pinus is among the best‐studied group of exotic species and one that has been widely introduced world‐wide, so we were able to obtain data on invasive and non‐invasive introductions (i.e. unsuccessful invasions; areas where after many decades of self‐sowing seeds there is no invasion). Location World‐wide. Methods We developed species distribution models for native ranges using a maximum entropy algorithm and projected them across the globe. We tested whether climate‐based models were able to predict both invasive and non‐invasive introductions. Results Appropriate climatic conditions seem to be required for these long‐lived species to invade because climates accurately predicted invasions. However, climate matching is necessary, but not sufficient to predict the fate of an introduction because most non‐invasive introductions were predicted to have triggered an invasion. Main conclusions Other factors, possibly including biotic components, may be the key to explaining why some introductions do not become invasions, because many areas where Pinus is not invading were predicted to be suitable for invasion based solely on climate.     

Quandaries of a decade‐long restoration experiment trying to reduce invasive species: beat them,join them,give up,or start over?

We evaluate the outcomes and consequences of a decade‐long restoration project in a Hawaiian lowland wet forest as they relate to long‐term management actions. Our initial study was designed both to promote native biodiversity and to develop knowledge that would enable land management agencies to restore invaded forests. Our premise of success followed the prevalent perception that short‐term management, such as removal of invasive species, ideally translates into long‐term and sustainable restoration. We were therefore disappointed and perhaps discouraged in our results—little recovery of native biodiversity despite ongoing and labor‐intensive management. Not only did we fail to return the invaded forest to a native‐dominated system but also our efforts lead to recruitment of new non‐native species assemblages. The sobering truth of many restoration projects in Hawaii and elsewhere is that we can never completely walk away and “consider the job finished,” or we have to accept that some ecosystems cannot be returned to an all‐native state. Essentially, costs of restoration may outweigh the accomplishment. This setback gave us an opportunity to reconsider and modify our initial approach. By starting over with a new direction using both native and non‐invasive but non‐native species, we have adopted a new philosophy of “join them.” In our revision, we changed the players in the game by following invasive species removal with outplantings of native and non‐invasive non‐native species that will functionally fill missing roles in the ecosystem. We link social interest in the new experiment to changing attitudes about naturalness.