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.     

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.

     

Invasive aliens on tropical East Asian islands

Tropical East Asia (TEA) has numerous islands, both continental and oceanic. This study uses information on invasive aliens in terrestrial habitats on these islands to test the generality of the continental-oceanic contrast in invasibility, assess the conservation impacts of invasive species, and suggest ways to mitigate these. The continental islands of Hong Kong and Singapore are worst-case scenarios for continental invasibility and alien species often dominate in chronically disturbed sites, but very few have successfully invaded closed forests, with the exception of birds in Hong Kong. On other, less densely populated, continental islands, closed-canopy forests appear to resist invasions by all taxa, with few known exceptions. Forests on oceanic islands isolated by <100 km during the last glacial maximum appear no more susceptible to plant and invertebrate invasions than those on continental islands, but invasions by mammals are widespread. Snake invasions may be under-recognized. The remote oceanic Ogasawara (Bonin) Islands, >1000 km from the nearest continent, have a native biota of largely tropical East Asian origin and are suffering from alien forest invasions across the taxonomic spectrum. These patterns of invasibility are consistent with the idea that alien invasion is facilitated by the absence of native species in the same functional group. Alien invasives are not yet a major conservation problem in TEA, except on remote islands, but their dominance on disturbed sites may slow or prevent recovery of native biodiversity. Strict quarantine is impractical in TEA, although some major introduction routes could be blocked. Management efforts should focus on early recognition and immediate control of potential problem species.     

Do alien plants on Mediterranean islands tend to invade different niches from native species?

In order to understand invasions, it is important to know how alien species exploit opportunities in unfamiliar ecosystems. For example, are aliens concentrated in niches under-exploited by native communities, or widely distributed across the ecological spectrum? To explore this question, we compared the niches occupied by 394 naturalized alien plants with a representative sample from the native flora of Mediterranean islands. When niche structure was described by a functional group categorization, the distribution of native and alien species was remarkably similar, although “succulent shrubs” and “trees with specialized animal pollination mechanisms” were under-represented in the native species pool. When niche structure was described by Grime’s CSR strategy, the positioning of aliens and natives differed more strongly. Stress-tolerance was much rarer amongst the aliens, and a competitive strategy was more prevalent at the habitat level. This pattern is similar to previous findings in temperate Europe, although in those regions it closely reflects patterns of native diversity. Stressed environments are much more dominant in the Mediterranean. We discuss a number of factors which may contribute to this difference, e.g., competitive and ruderal niches are often associated with anthropogenic habitats, and their high invasibility may be due partly to introduction patterns rather than to a greater efficiency of aliens at exploiting them. Thus far, the reasons for invasion success amongst introduced species have proved difficult to unravel. Despite some differences, our evidence suggests that alien species naturalize across a wide range of niches. Given that their ecologies therefore vary greatly, one may ask why such species should be expected to share predictable traits at all?     

Species pool size and invasibility of island communities: a null model of sampling effects

The success of alien species on oceanic islands is considered to be one of the classic observed patterns in ecology. Explanations for this pattern are based on lower species richness on islands and the lower resistance of species‐poor communities to invaders, but this argument needs re‐examination. The important difference between islands and mainland is in the size of species pools, not in local species richness; invasibility of islands should therefore be addressed in terms of differences in species pools. Here I examine whether differences in species pools can affect invasibility in a lottery model with pools of identical native and exotic species. While in a neutral model with all species identical, invasibility does not depend on the species pool, a model with non‐zero variation in population growth rates predicts higher invasibility of communities of smaller pools. This is because of species sampling; drawing species from larger pools increases the probability that an assemblage will include fast growing species. Such assemblages are more likely to exclude random invaders. This constitutes a mechanism through which smaller species pools (such as those of isolated islands) can directly underlie differences in invasibility.     

Do leaf margins of the temperate forest flora of southern South America reflect a warmer past?

Aim   Our aims were: (1) to characterize the linear relationship between the proportion of woody dicotyledonous species with entire-margined leaves ( E ) and mean annual temperature (MAT) from a southern temperate flora that still harbours many lineages that originated under warmer climates; (2) to compare this relationship with those developed from floras of different regions of the world; and (3) to contrast temperature predictions based on leaf margins of the native southern flora versus the naturalized alien flora, mostly of boreal origin.
Location   The temperate forest of southern South America (TFSA).
Methods   At each 1° latitudinal band, we estimated E based on species latitudinal ranges and MAT from both an isotherm map and a global temperature grid. We also calculated E from five local floras located between 40 and 43° S, and from the naturalized alien flora of Nahuel Huapi National Park in southern Argentina.
Results   We found a close relationship between E and MAT for the TFSA. Equations developed from floras of the Northern Hemisphere overestimated extant temperatures of this biome by 6–10 °C at both geographical and local spatial scales. On the other hand, MAT predictions from leaf margins of the alien flora were similar to the actual MAT. A published regression between E and MAT from tropical South America was remarkably similar to the one we estimated from the TFSA. This tropical equation predicted accurately the temperatures observed for this temperate biome based on leaf margins of the native flora.
Main conclusions   Despite massive plant extinction due to environmental cooling and biogeographical isolation during the Tertiary, leaf-margin analysis reveals that the flora of the TFSA still reflects its original development under the warmer conditions of western Gondwana and its past connections with low-latitude forest floras of tropical South America.     

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.     

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.     

How general are global trends in biotic homogenization? Floristic tracking in Chile,South America

Aim   To quantify the occurrence of floristic change in the vascular flora of Chile. We test whether continental areas have experienced floristic modification leading to either homogenization, differentiation or tracking.
Location   Continental Chile.
Methods   On the basis of the geographical distribution of native (1806 species) and naturalized plants (552 species) in continental Chile, we quantified change between two floristic stages: (1) pre-European flora, including native extant and extinct species; and (2) current flora, including native and naturalized species, but excluding extinct plants. We compared changes in compositional similarity (calculated by Jaccard's index, Δ J ) between pairs of regions, and similarity decay with respect to geographical distance. Additionally, by means of Whittaker's index, we examined species turnover, distinguishing between native and naturalized plants.
Results   Between floristic stages (pre-European vs. current flora) no significant changes in floristic similarity were noted at national or regional scales. Similarity decay showed no statistical differences between pre-European and current flora. Analysing patterns of geographical turnover, we found that species turnover of naturalized plants over their geographical range is similar to that of native plants.
Conclusions   The composition of the continental flora of Chile does not show significant modifications in similarity patterns after considering naturalized species, thus indicating floristic tracking. The causes of this phenomenon may be related to the current geographical distribution of naturalized plants, which closely parallels that of native plants. Our results differ from those obtained in Northern Hemisphere continents, thus indicating that trends of biotic change may differ between hemispheres.     

Are island plant communities more invaded than their mainland counterparts?

Questions: Are island vegetation communities more invaded than their mainland counterparts? Is this pattern consistent among community types? Location: The coastal provinces of Catalonia and the para‐oceanic Balearic Islands, both in NE Spain. These islands were connected to the continent more than 5.35 million years ago and are now located <200 km from the coast. Methods: We compiled a database of almost 3000 phytosociological relevés from the Balearic Islands and Catalonia and compared the level of invasion by alien plants in island versus mainland communities. Twenty distinct plant community types were compared between island and mainland counterparts. Results: The percentage of plots with alien species, number, percentage and cover percentage of alien species per plot was greater in Catalonia than in the Balearic Islands in most communities. Overall, across communities, more alien species were found in the mainland (53) compared to the islands (only nine). Despite these differences, patterns of the level of invasion in communities were highly consistent between the islands and mainland. The most invaded communities were ruderal and riparian. Main conclusion: Our results indicate that para‐oceanic island communities such as the Balearic Islands are less invaded than their mainland counterparts. This difference reflects a smaller regional alien species pool in the Balearic Islands than in the adjacent mainland, probably due to differences in landscape heterogeneity and propagule pressure.