Traits associated with naturalization in <Emphasis Type="Italic">Anolis</Emphasis> lizards: comparison of morphological,distributional, anthropogenic,and phylogenetic models

The worldwide spread of invasive species affects native biodiversity and causes economic loss, but also allows better understanding of historical biogeographic patterns. Prediction of likely invaders facilitates economic and conservation decisions and gives insight into characteristics that have allowed natural colonization over evolutionary time. However, it is not clear what types of characters best predict naturalization or even whether naturalization is predictable at all. Squamate reptiles have been understudied subjects for invasion biology. Lizards of the genus Anolis have been highly successful colonizers both recently and over evolutionary time. Nineteen of the approximately 350 described species of Anolis have established naturalized populations. We constructed models of naturalization using morphological, distributional, anthropogenic, and phylogenetic characters and compared these single character class models to each other and to a composite model incorporating all four classes. We show that (1) each class of variables significantly predicts invasion, (2) a composite model significantly outperforms each of the submodels, and (3) the final composite model displays extraordinary ability to objectively identify naturalized species of Anolis.     

Searching for phylogenetic pattern in biological invasions

It has been suggested that alien species with close indigenous relatives in the introduced range may have reduced chances of successful establishment and invasion (Darwin's naturalization hypothesis). Studies trying to test this have in fact been addressing four different hypotheses, and the same data can support some while rejecting others. In this paper, we argue that the phylogenetic pattern will change depending on the spatial and phylogenetic scales considered. Expectations and observations from invasion biology and the study of natural communities are that at the spatial scale relevant to competitive interactions, closely related species will be spatially separated, whereas at the regional scale, species in the same genera or families will tend to co-occur more often than by chance. We also argue that patterns in the relatedness of indigenous and naturalized plants are dependent on the continental/island setting, spatial occupancy levels, and on the group of organisms under scrutiny. Understanding how these factors create a phylogenetic pattern in invasions will help us predict which groups are more likely to invade where, and should contribute to general ecological theory.     

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

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

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.     

Convergent exaptation and adaptation in solitary island lizards

Independent evolutionary lineages often display similar characteristics in comparable environments. Three kinds of historical hypotheses could explain this convergence. The first is adaptive and evolutionary: nonrandom patterns may result from analogous evolutionary responses to shared conditions. The second explanation is exaptive and ecological: species may be filtered by their suitability for a particular type of environment. The third potential explanation is a null hypothesis of random colonization from a historically nonrandom source pool. Here we demonstrate that both exaptation and adaptation have produced convergent similarity in different size-related characters of solitary island lizards. Large sexual size dimorphism results from adaptive response to solitary existence; uniform, intermediate size results from ecological filtering of potential colonizers. These results demonstrate the existence of deterministic exaptive convergence and suggest that convergent phenomena may require historical explanations that are ecological as well as evolutionary.     

Ethnicity and naturalization

This article examines whether or not ethnicity has an independent effect on the likelihood of immigrant naturalization using the Public Use Microdata Sample [PUMS] data from the 1980 US Census. Ethnic differences in the propensity to become naturalized US citizens were analysed among four panethnic groups and across thirty‐three major ethnic groups. The results point to the continuing significance of ethnicity in the naturalization process. However, the effect of ethnicity is not as strong as the effects of other structural factors. Three hypotheses that attempt to explain ethnic differences in the propensity for naturalization were also tested. The evidence lends strong support to the forced self‐protection hypothesis, but it provides no support for the discrimination hypothesis and the cultural differences hypothesis.     

Hybridization-prone plant families do not generate more invasive species

Many plant taxa are both hybrid-derived and invasive, suggesting a causal connection. However, given that hybridization is not rare in plants, we should expect some fraction of invasive taxa to be hybrids, even in the absence of an underlying causal relationship. Here, we test the hypothesis that hybridization leads to invasiveness by asking whether the number of hybrids and the numbers of naturalized, weedy, and invasive taxa are correlated across 256 vascular plant families. Data were derived from six regional floras and three global databases listing weeds and invasives. To account for phylogenetic nonindependence, we combined a supertree analysis with phylogenetically independent contrasts. After correcting for family size and phylogeny, we conclude that vascular plant families with a higher propensity for hybridization are not more likely to produce more naturalized, weedy, or invasive species than families less prone to hybridization. Instead, hybridization-prone families were in some cases associated with fewer naturalized species and invaders. We present two hypotheses for these patterns, one based on Levin’s (Syst Bot 31:8–12, 2006) ideas on reproductive interference and another based on Darwin’s naturalization hypothesis. While these results do not preclude the possibility that hybridization generates weedy and invasive taxa with some frequency, they do suggest that the signal from the hybridization-invasion process may be relatively weak and easily obscured by other processes governing plant invasions.     

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.     

Naturalization of ornamental plant species in public green spaces and private gardens

Ornamental horticulture is the most important pathway for alien plant introductions worldwide, and consequently, invasive spread of introduced plants often begins in urban areas. Although most introduced ornamental garden-plant species are locally not naturalized yet, many of them have shown invasion potential elsewhere in the world, and might naturalize when climate changes. We inventoried the planted flora of 50 public and 61 private gardens in Radolfzell, a small city in southern Germany, to investigate whether local naturalization success of garden plants is associated with their current planting frequency, climatic suitability (as assessed with climatic niche modelling) and known naturalization status somewhere in the world. We identified 954 introduced garden-plant species, of which 48 are already naturalized in Radolfzell and 120 in other parts of Germany. All currently naturalized garden plants in Radolfzell have a climatic suitability probability of ≥ 0.75 and are naturalized in ≥ 13 out of 843 regions globally. These values are significantly higher than those of garden plants that have not become locally naturalized yet. Current planting frequencies, however, were not related to current naturalization success. Using the identified local naturalization thresholds of climatic suitability and global naturalization frequency, and climate projections for the years 2050 and 2070, we identified 45 garden-plant species that are currently not naturalized in Radolfzell but are likely to become so in the future. Although our approach cannot replace a full risk assessment, it is well-suited and applicable as one element of a screening or horizon scanning-type approach.     

外来种-本地种亲缘关系对外来植物归化和入侵的影响

达尔文归化难题描述了外来种-本地种亲缘关系促进(预适应假说)或阻止(达尔文归化假说)外来种成功入侵的悖论。目前, 在中国仍缺少针对达尔文归化难题的研究。为系统研究外来种-本地种亲缘关系对中国外来植物入侵的影响, 该文利用线性混合效应模型从省级、市级和群落3个空间尺度以及归化、扩散和入侵3个阶段探究了外来种-本地种谱系距离和外来植物表现的关系。结果表明: 在省级和市级(区域)尺度上, 与本地种亲缘关系较近的外来植物更有可能在当地归化和扩散, 符合预适应假说的预期; 而在群落(局域)尺度上, 外来种-本地种亲缘距离与外来种是否在群落中成功定居及其入侵程度无关。该研究结果表明与本地区系亲缘较近的外来种和本地种的竞争并不强烈, 却能较好地适应本地气候环境而具有更强的归化和入侵潜力。因此, 在今后的外来植物管理和治理中需要尤其重视与本地区系亲缘关系较近的外来植物。     

Effects of exotic-native species relationship on naturalization and invasion of exotic plant species北大核心CSCD

Aims: Darwin's naturalization conundrum describes the paradox that the relationship of exotic species to native residents could either promote or hinder invasion success through opposing mechanisms: niche pre-adaptation or competitive interactions. Previous Darwin's naturalization studies have showed invasion success could vary at stages, sites, and spatial and phylogenetic scales. Our objective was to assess the effects of exotic-native species relationship on invasion process of exotic plant species in China, where related research is still lacking. Methods: Generalized linear mixed models were used to examine relationship between exotic-native species relationship and performance of exotic species at different spatial scale (provincial, municipal and community) and invasion stages (naturalization, dispersal and invasion). At community scale, we measured environmental factors of communities we investigated to control the effect of habitat heterogeneity among them. Important findings: At the provincial and municipal scales, exotic species closely related to native flora were more likely to be naturalized and distributed, which is more consistent with the expectation of the pre-adaptation hypothesis. On the community scale, the exotic-native species relationship was not related to establishment and abundance of exotic species in the community. The results suggested that exotic species did not strongly compete with their close native relatives in communities, but were better adapted to areas where their close relatives had lived. Considering their high potential of naturalization and invasion, special attention should be paid to those exotic species that closely related to the native flora in the management of invasive species. © Editorial Office of Chinese Journal of Plant Ecology. All rights reserved.     

Effects of exotic-native species relationship on naturalization and invasion of exotic plant species

达尔文归化难题描述了外来种-本地种亲缘关系促进(预适应假说)或阻止(达尔文归化假说)外来种成功入侵的悖论。目前, 在中国仍缺少针对达尔文归化难题的研究。为系统研究外来种-本地种亲缘关系对中国外来植物入侵的影响, 该文利用线性混合效应模型从省级、市级和群落3个空间尺度以及归化、扩散和入侵3个阶段探究了外来种-本地种谱系距离和外来植物表现的关系。结果表明: 在省级和市级(区域)尺度上, 与本地种亲缘关系较近的外来植物更有可能在当地归化和扩散, 符合预适应假说的预期; 而在群落(局域)尺度上, 外来种-本地种亲缘距离与外来种是否在群落中成功定居及其入侵程度无关。该研究结果表明与本地区系亲缘较近的外来种和本地种的竞争并不强烈, 却能较好地适应本地气候环境而具有更强的归化和入侵潜力。因此, 在今后的外来植物管理和治理中需要尤其重视与本地区系亲缘关系较近的外来植物。     

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.     

Darwin's naturalization hypothesis: scale matters in coastal plant communities

Darwin proposed two seemingly contradictory hypotheses for a better understanding of biological invasions. Strong relatedness of invaders to native communities as an indication of niche overlap could promote naturalization because of appropriate niche adaptation, but could also hamper naturalization because of negative interactions with native species (‘Darwin's naturalization hypothesis’). Although these hypotheses provide clear and opposing predictions for expected patterns of species relatedness in invaded communities, so far no study has been able to clearly disentangle the underlying mechanisms. We hypothesize that conflicting past results are mainly due to the neglected role of spatial resolution of the community sampling. In this study, we corroborate both of Darwin's expectations by using phylogenetic relatedness as a measure of niche overlap and by testing the effects of sampling resolution in highly invaded coastal plant communities. At spatial resolutions fine enough to detect signatures of biotic interactions, we find that most invaders are less related to their nearest relative in invaded plant communities than expected by chance (phylogenetic overdispersion). Yet at coarser spatial resolutions, native assemblages become more invasible for closely‐related species as a consequence of habitat filtering (phylogenetic clustering). Recognition of the importance of the spatial resolution at which communities are studied allows apparently contrasting theoretical and empirical results to be reconciled. Our study opens new perspectives on how to better detect, differentiate and understand the impact of negative biotic interactions and habitat filtering on the ability of invaders to establish in native communities.     

Limiting similarity and Darwin’s naturalization hypothesis: understanding the drivers of biotic resistance against invasive plant species

Several hypotheses have been proposed to explain biotic resistance of a recipient plant community based on reduced niche opportunities for invasive alien plant species. The limiting similarity hypothesis predicts that invasive species are less likely to establish in communities of species holding similar functional traits. Likewise, Darwin’s naturalization hypothesis states that invasive species closely related to the native community would be less successful. We tested both using the invasive alien Ambrosia artemisiifolia L. and Solidago gigantea Aiton, and grassland species used for ecological restoration in central Europe. We classified all plant species into groups based on functional traits obtained from trait databases and calculated the phylogenetic distance among them. In a greenhouse experiment, we submitted the two invasive species at two propagule pressures to competition with communities of ten native species from the same functional group. In another experiment, they were submitted to pairwise competition with native species selected from each functional group. At the community level, highest suppression for both invasive species was observed at low propagule pressure and not explained by similarity in functional traits. Moreover, suppression decreased asymptotically with increasing phylogenetic distance to species of the native community. When submitted to pairwise competition, suppression for both invasive species was also better explained by phylogenetic distance. Overall, our results support Darwin’s naturalization hypothesis but not the limiting similarity hypothesis based on the selected traits. Biotic resistance of native communities against invasive species at an early stage of establishment is enhanced by competitive traits and phylogenetic relatedness.     

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.     

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.

     

Does predation select for or against avian coloniality? A comparative analysis

Some studies have supported predation as a selective pressure contributing to the evolution of coloniality. However, evidence also exists that colonies attract predators, selecting against colonial breeding. Using comparative analyses, we tested the reduced predation hypothesis that individuals aggregate into colonies for protection, and the opposite hypothesis, that breeding aggregations increase predation risk. We used locational and physical characteristics of nests to estimate levels of species' vulnerability to predation. We analysed the Ciconiiformes, a large avian order with the highest prevalence of coloniality, using Pagel's general method of comparative analysis for discrete variables. A common requirement of both hypotheses, that there is correlated evolution between coloniality and vulnerability to predation, was fulfilled in our data set of 363 species. The main predictions of the reduced predation hypothesis were not supported, namely that (1) solitary/vulnerable species are more prone to become colonial than solitary/protected species and (2) colonial/protected species are more likely to evolve towards vulnerability than solitary/protected species. In contrast, the main predictions of the increased predation hypothesis were supported, namely that colonial/vulnerable species are more prone (1) to become protected than solitary/vulnerable species and/or (2) to become solitary than colonial/protected species. This suggests that the colonial/vulnerable state is especially exposed to predation as coloniality may often attract predators rather than provide safety.     

'Nasty neighbours' rather than 'dear enemies' in a social carnivore

Territorial animals typically respond less aggressively to neighbours than to strangers. This 'dear enemy effect' has been explained by differing familiarity or by different threat levels posed by neighbours and strangers. In most species, both the familiarity and the threat-level hypotheses predict a stronger response to strangers than to neighbours. In contrast, the threat-level hypothesis predicts a stronger response to neighbours than to strangers in species with intense competition between neighbours and with residents outnumbering strangers, as commonly found in social mammals such as the banded mongoose (Mungos mungo). The familiarity hypothesis predicts reduced aggression towards neighbours also in these species. We exposed free-living banded mongoose groups to translocated scent marks of neighbouring groups and strangers. Groups vocalized more and inspected more samples in response to olfactory cues of the neighbours than to the strangers. Our results support the threat-level hypothesis and contradict the familiarity hypothesis. We suggest that increased aggression towards neighbours is more common in social species with intense competition between neighbours, as opposed to reduced aggression towards neighbours typical for most solitary species.