Naturalization and invasion of alien plants: concepts and definitions

Abstract.  Much confusion exists in the English-language literature on plant invasions concerning the terms 'naturalized' and 'invasive' and their associated concepts. Several authors have used these terms in proposing schemes for conceptualizing the sequence of events from introduction to invasion, but often imprecisely, erroneously or in contradictory ways. This greatly complicates the formulation of robust generalizations in invasion ecology.
Based on an extensive and critical survey of the literature we defined a minimum set of key terms related to a graphic scheme which conceptualizes the naturalization/invasion process. Introduction means that the plant (or its propagule) has been transported by humans across a major geographical barrier. Naturalization starts when abiotic and biotic barriers to survival are surmounted and when various barriers to regular reproduction are overcome. Invasion further requires that introduced plants produce reproductive offspring in areas distant from sites of introduction (approximate scales: > 100 m over < 50 years for taxa spreading by seeds and other propagules; > 6 m/3 years for taxa spreading by roots, rhizomes, stolons or creeping stems). Taxa that can cope with the abiotic environment and biota in the general area may invade disturbed, seminatural communities. Invasion of successionally mature, undisturbed communities usually requires that the alien taxon overcomes a different category of barriers.
We propose that the term 'invasive' should be used without any inference to environmental or economic impact. Terms like 'pests' and 'weeds' are suitable labels for the 50–80% of invaders that have harmful effects. About 10% of invasive plants that change the character, condition, form, or nature of ecosystems over substantial areas may be termed 'transformers'.     

Herbivores and the success of exotic plants: a phylogenetically controlled experiment

In a field experiment with 30 locally occurring old‐field plant species grown in a common garden, we found that non‐native plants suffer levels of attack (leaf herbivory) equal to or greater than levels suffered by congeneric native plants. This phylogenetically controlled analysis is in striking contrast to the recent findings from surveys of exotic organisms, and suggests that even if ‘enemy release’ does accompany the invasion process, this may not be an important mechanism of invasion, particularly for plants with close relatives in the recipient flora.     

Undaria pinnatifida (Laminariales,Phaeophyta) 12 years after its introduction into the Atlantic Ocean

Since its first introduction into the Atlantic for farming purposes in 1983, the brown alga Undaria pinnatifida has been found at several sites along the European coasts. The present study deals with an ecological experiment that was performed at the island of Ushant (Brittany, France) in order to check the ability of Undaria to recruit to a rocky substratum in situ. On the one hand, the authors were unsuccessful in getting Undaria to colonize a previously denuded area of shore, which was subsequently colonized by other native species, a result that may speak for a competition between Undaria and the local kelps. On the other hand, evidence is presented that Undaria pinnatifida has become a member of the Atlantic flora for good. The sporophytes can be found growing from low water mark of neap tide (+1.5 m) down to at least –15 m below lower water mark of spring tide. Its propagation is thought to occur in at least two ways: either step by step from cultivation sites along the coasts, or from the hulls of ships sailing from harbour to harbour. The preference of Undaria for settlement is on artificial structures.     

Invasion success of alien plants: do habitat affinities in the native distribution range matter?

Aim To assess how habitat affinities in the native distribution range influence the invasion success of 282 central European neophytes (alien plants introduced after ad 1500). Location Czech Republic. Methods Classification trees were used to determine which native habitats donate the most alien species, the correspondence between habitats occupied by species in their native and invaded distribution ranges, and invasion success of species originating from different habitats. Results The species most likely to naturalize in Central Europe are those associated with thermophile woodland fringes in their native range (81%), cultivated areas of gardens and parks (75%) and broad‐leaved deciduous woodlands (72%). The largest proportions of invasive species recruit from those that occur on riverine terraces and eroded slopes, or grow in both deciduous woodland and riverine scrub. When the relative role of habitats in the native range is assessed as a determinant of the probability that a species will become invasive in concert with other factors (the species’ residence time, life history, region of origin), the direct effect of habitat is negligible. However, the effect of native habitats on patterns of invasions observed in central Europe is manifested by large differences in the numbers of species they supply to the invaded region. More than 50 neophytes were recruited from each of the following habitats: dry grasslands, ruderal habitats, deciduous woodland, inland cliffs, rock pavements and outcrops, and tall‐herb fringes and meadows. Main conclusions Casual species recruit from a wider range of habitats in their native range than they occupy in the invaded range; naturalized but not invasive species inhabit a comparable spectrum of habitats in both ranges, and successful invaders occupy a wider range of habitats in the invaded than in the native range. This supports the idea that the invasive phase of the process is associated with changes in biological features that allow for extension of the spectrum of habitats invaded.     

Biological invasions and scientific objectivity: Reply to Cassey et al. (2005)

Abstract We disagree with the assertion that recent human‐caused invasions differ substantially from historic natural invasions in their magnitudes and impacts on ecological processes. The position that exotic species are inherently ‘bad’ and should be eradicated is an ethical judgement, usually based on the naturalist fallacy or xenophobic prejudice; it is not a scientific judgement. The role of scientists in studying invasive species should be to gather, interpret and communicate information as accurately and objectively as possible.     

Revegetation following Soil Disturbance and Invasion in a Californian Meadow: a 10-year History of Recovery

Disturbance is necessary for the regeneration of many native plant species, but can also facilitate biological invasions. As a result, disturbance can play complex roles in vulnerable habitats such as remnant Californian perennial grasslands. To investigate these conflicts, plots in a northern Californian coastal grassland were experimentally disturbed in the winter of 1990–1991; these plots differed in the area and intensity (depth) of the soil disturbance applied. When these plots were revisited after 10 growing seasons, patterns of revegetation differed significantly from those observed early in recolonization (0–3 years). At the earlier samplings, exotic annual grasses rapidly increased in most disturbance types. After 10 years, these exotic annuals had retreated from the depth experiment, which had recovered to a vegetation dominated by native perennials in all but the most severely disturbed plots. In contrast, although differences between control and disturbed plots also disappeared in the area experiment, the average abundance of aliens did not decline substantially relative to 1993 levels, especially in larger disturbances. Nonetheless, populations of aliens remained small compared to the peak populations in the depth experiment, probably reflecting wetter soils at the site used for the area experiment. These results differ from those of other recent studies of soil disturbance in coastal Californian ecosystems, which indicate disturbance may result in the permanent replacement of native perennial vegetation by dense populations of exotic annual grasses. This difference may reflect the high resilience of northern coastal grasslands as well as the scale of disturbances considered by different studies.     

Fronts,jumps and secondary introductions suggested as different invasion patterns in marine species,with an increase in spread rates over time

Not all introduced (invasive) species in a region will spread from a single point of introduction. Long-distance dispersal or further introductions can obscure the pattern of spread, but the regional importance of such processes is difficult to gauge. These difficulties are further compounded when information on the multiple scale process of invasive species range expansion is reduced to one-dimensional estimates of spread (e.g. km yr⁻¹). We therefore compared the results of two different metrics of range expansion: maximum linear rate of spread and accumulation of occupied grid squares (50 × 50 km) over time. An analysis of records for 54 species of introduced marine macrophytes in the Mediterranean and northeast Atlantic revealed cases where the invasion process was probably missed (e.g. Atlantic Bonnemaisonia hamifera) and suggested cases of secondary introductions or erratic jump dispersal (Dasysiphonia sp. and Womersleyella setacea). A majority of species analysed showed evidence for an accumulation of invaded sites without a clear invasion front. Estimates of spread rate are increasing for more recent introductions. The increase is greater than can be accounted for by temporally varying search effort and implies a historical increase in vector efficiency and/or a decreased resistance of native communities to invasion.     

Survival rates in the Czech Republic of introduced plants known as wool aliens

The paper analyses an unintentional experiment conducted 40 years ago with species termed wool aliens which were observed in employee garden allotments of a wool-processing factory in Brno, Czech Republic. The waste from processing was used as a garden fertilizer, and alien species were recorded by Dvořák and Kühn (1966) Zavlečené rostliny na pozemcích přádelny vlny ‘Mosilana’ n.p. v Brně. Preslia 38: 327–332. The survival of these species was assessed by their occurrence in the current alien flora of the Czech Republic. Of the 56 species introduced in the 1960s and classified as wool aliens, 18 are still present in the Czech alien flora. The remaining 38 species are considered extinct, i.e. alien plants not capable of long-term survival in the country. The resulting survival rate of wool aliens is 22.1%. Three species on the list (5.4%) were clearly capable of naturalization. Compared to the other alien flora of the Czech Republic, wool aliens were disproportionally introduced from Australia, Africa and South and Central America. Species from Europe and Africa tended to survive the best. Grasses were most strongly represented, contributing 62.5% to wool alien species but only 7.9% to the total Czech neophyte flora. All surviving species were annuals, whereas none of the perennials survived; the survival rate in annuals was 37.5%. Survival of species introduced with wool was significantly comparable to their performance in the UK and in Central Europe, indicating that survival is not determined by mere chance and that the same species tend to be successful elsewhere in Europe. Maximum height and the month of first flowering were not significantly related to the species survival. The role of specific events in the enrichment of alien floras is discussed.     

Alien plants in central Australia

The central Australian flora currently contains 78 alien species, of which 17 are predominantly summer-growing, 19 are southern winter-growing species reaching roadsides, stockyards and watercourses in the southern Northern Territory and 42 are at present confined to gardens in Alice Springs. A further 11 species reach the far north of South Australia or die far southwest of Queensland but not the Northern Territory. The central Australian alien flora may be classified by growing season and drought-tolerance, or broadly categorized on the basis of habitat and dispersal agent into 'garden weeds', 'tourist weeds' and 'stock weeds'. Present ecological knowledge of central Australian aliens is limited, but indicates that whilst some will probably remain confined to better-watered habitats, many are likely to spread into the arid areas, becoming obvious only after a succession of wet years. The central Australian alien flora is increasing at present and this increase is expected to continue.