Climatic stability and geological history shape global centers of neo- and paleoendemism in seed plants.

Assessing the distribution of geographically restricted and evolutionarily unique species and their underlying drivers is key to understanding biogeographical processes and critical for global conservation prioritization. Here, we quantified the geographic distribution and drivers of phylogenetic endemism for ~320,000 seed plants worldwide and identified centers and drivers of evolutionarily young (neoendemism) and evolutionarily old endemism (paleoendemism). Tropical and subtropical islands as well as tropical mountain regions displayed the world's highest phylogenetic endemism. Most tropical rainforest regions emerged as centers of paleoendemism, while most Mediterranean-climate regions showed high neoendemism. Centers where high neo- and paleoendemism coincide emerged on some oceanic and continental fragment islands, in Mediterranean-climate regions and parts of the Irano-Turanian floristic region. Global variation in phylogenetic endemism was well explained by a combination of past and present environmental factors (79.8 to 87.7% of variance explained) and most strongly related to environmental heterogeneity. Also, warm and wet climates, geographic isolation, and long-term climatic stability emerged as key drivers of phylogenetic endemism. Neo- and paleoendemism were jointly explained by climatic and geological history. Long-term climatic stability promoted the persistence of paleoendemics, while the isolation of oceanic islands and their unique geological histories promoted neoendemism. Mountainous regions promoted both neo- and paleoendemism, reflecting both diversification and persistence over time. Our study provides insights into the evolutionary underpinnings of biogeographical patterns in seed plants and identifies the areas on Earth with the highest evolutionary and biogeographical uniqueness-key information for setting global conservation priorities.

Regional invasion history and land use shape the prevalence of non-native species in local assemblages.

The ecological impact of non-native species arises from their establishment in local assemblages. However, the rates of non-native spread in new regions and their determinants have not been comprehensively studied. Here, we combined global databases documenting the occurrence of non-native species and residence of non-native birds, mammals, and vascular plants at regional and local scales to describe how the likelihood of non-native occurrence and their proportion in local assemblages relate with their residence time and levels of human usage in different ecosystems. Our findings reveal that local non-native occurrence generally increases with residence time. Colonization is most rapid in croplands and urban areas, while it is slower and variable in natural or semi-natural ecosystems. Notably, non-native occurrence continues to rise even 200 years after introduction, especially for birds and vascular plants, and in other land-use types rather than croplands and urban areas. The impact of residence time on non-native proportions is significant only for mammals. We conclude that the continental exchange of biotas requires considerable time for effects to manifest at the local scale across taxa and land-use types. The unpredictability of future impacts, implied by the slow spread of non-native species, strengthens the call for stronger regulations on the exchange of non-native species to reduce the long-lasting invasion debt looming on ecosystems' future.

Small genome size and variation in ploidy levels support the naturalization of vascular plants but constrain their invasive spread.

Karyological characteristics are among the traits underpinning the invasion success of vascular plants. Using 11 049 species, we tested the effects of genome size and ploidy levels on plant naturalization (species forming self-sustaining populations where they are not native) and invasion (naturalized species spreading rapidly and having environmental impact). The probability that a species naturalized anywhere in the world decreased with increasing monoploid genome size (DNA content of a single chromosome set). Naturalized or invasive species with intermediate monoploid genomes were reported from many regions, but those with either small or large genomes occurred in fewer regions. By contrast, large holoploid genome sizes (DNA content of the unreplicated gametic nucleus) constrained naturalization but favoured invasion. We suggest that a small genome is an advantage during naturalization, being linked to traits favouring adaptation to local conditions, but for invasive spread, traits associated with a large holoploid genome, where the impact of polyploidy may act, facilitate long-distance dispersal and competition with other species.

The big four of plant taxonomy - a comparison of global checklists of vascular plant names.

Taxonomic checklists used to verify published plant names and identify synonyms are a cornerstone of biological research. Four global authoritative checklists for vascular plants exist: Leipzig Catalogue of Vascular Plants, World Checklist of Vascular Plants, World Flora Online (successor of The Plant List, TPL), and WorldPlants. We compared these four checklists in terms of size and differences across taxa. We matched taxon names of these checklists and TPL against each other, identified differences across checklists, and evaluated the consistency of accepted names linked to individual taxon names. We assessed geographic and phylogenetic patterns of variance. All checklists differed strongly compared with TPL and provided identical information on c. 60% of plant names. Geographically, differences in checklists increased from low to high latitudes. Phylogenetically, we detected strong variability across families. A comparison of name-matching performance on taxon names submitted to the functional trait database TRY, and a check of completeness of accepted names evaluated against an independent, expert-curated checklist of the family Meliaceae, showed a similar performance across checklists. This study raises awareness on the differences in data and approach across these checklists potentially impacting analyses. We propose ideas on the way forward exploring synergies and harmonizing the four global checklists.

Global models and predictions of plant diversity based on advanced machine learning techniques.

Despite the paramount role of plant diversity for ecosystem functioning, biogeochemical cycles, and human welfare, knowledge of its global distribution is still incomplete, hampering basic research and biodiversity conservation. Here, we used machine learning (random forests, extreme gradient boosting, and neural networks) and conventional statistical methods (generalized linear models and generalized additive models) to test environment-related hypotheses of broad-scale vascular plant diversity gradients and to model and predict species richness and phylogenetic richness worldwide. To this end, we used 830 regional plant inventories including c. 300 000 species and predictors of past and present environmental conditions. Machine learning showed a superior performance, explaining up to 80.9% of species richness and 83.3% of phylogenetic richness, illustrating the great potential of such techniques for disentangling complex and interacting associations between the environment and plant diversity. Current climate and environmental heterogeneity emerged as the primary drivers, while past environmental conditions left only small but detectable imprints on plant diversity. Finally, we combined predictions from multiple modeling techniques (ensemble predictions) to reveal global patterns and centers of plant diversity at multiple resolutions down to 7774 km2 . Our predictive maps provide accurate estimates of global plant diversity available at grain sizes relevant for conservation and macroecology.

Persistent soil seed banks promote naturalisation and invasiveness in flowering plants.

With globalisation facilitating the movement of plants and seeds beyond the native range, preventing potentially harmful introductions requires knowledge of what drives the successful establishment and spread of alien plants. Here, we examined global-scale relationships between naturalisation success (incidence and extent) and invasiveness, soil seed bank properties (type and densities) and key species traits (seed mass, seed dormancy and life form) for 2350 species of angiosperms. Naturalisation and invasiveness were strongly associated with the ability to form persistent (vs. transient) seed banks but relatively weakly with seed bank densities and other traits. Our findings suggest that seed bank persistence is a trait that better captures the ability to become naturalised and invasive compared to seed traits more widely available in trait databases. Knowledge of seed persistence can contribute to our ability to predict global naturalisation and invasiveness and to identify potentially invasive flowering plants before they are introduced.

Role of diversification rates and evolutionary history as a driver of plant naturalization success.

Human introductions of species beyond their natural ranges and their subsequent establishment are defining features of global environmental change. However, naturalized plants are not uniformly distributed across phylogenetic lineages, with some families contributing disproportionately more to the global alien species pool than others. Additionally, lineages differ in diversification rates, and high diversification rates have been associated with characteristics that increase species naturalization success. Here, we investigate the role of diversification rates in explaining the naturalization success of angiosperm plant families. We use five global data sets that include native and alien plant species distribution, horticultural use of plants, and a time-calibrated angiosperm phylogeny. Using phylogenetic generalized linear mixed models, we analysed the effect of diversification rate, different geographical range measures, and horticultural use on the naturalization success of plant families. We show that a family's naturalization success is positively associated with its evolutionary history, native range size, and economic use. Investigating interactive effects of these predictors shows that native range size and geographic distribution additionally affect naturalization success. High diversification rates and large ranges increase naturalization success, especially of temperate families. We suggest this may result from lower ecological specialization in temperate families with large ranges, compared with tropical families with smaller ranges.

Disturbed habitats locally reduce the signal of deep evolutionary history in functional traits of plants.

The functioning of present ecosystems reflects deep evolutionary history of locally cooccurring species if their functional traits show high phylogenetic signal (PS). However, we do not understand what drives local PS. We hypothesize that local PS is high in undisturbed and stressful habitats, either due to ongoing local assembly of species that maintained ancestral traits, or to past evolutionary maintenance of ancestral traits within habitat species-pools, or to both. We quantified PS and diversity of 10 traits within 6704 local plant communities across 38 Dutch habitat types differing in disturbance or stress. Mean local PS varied 50-fold among habitat types, often independently of phylogenetic or trait diversity. Mean local PS decreased with disturbance but showed no consistent relationship to stress. Mean local PS exceeded species-pool PS, reflecting nonrandom subsampling from the pool. Disturbance or stress related more strongly to mean local than to species-pool PS. Disturbed habitats harbour species with evolutionary divergent trait values, probably driven by ongoing, local assembly of species: environmental fluctuations might maintain different trait values within lineages through an evolutionary storage effect. If functional traits do not reflect phylogeny, ecosystem functioning might not be contingent on the presence of particular lineages, and lineages might establish evolutionarily novel interactions.

Global exchange and accumulation of non-native plants.

All around the globe, humans have greatly altered the abiotic and biotic environment with ever-increasing speed. One defining feature of the Anthropocene epoch is the erosion of biogeographical barriers by human-mediated dispersal of species into new regions, where they can naturalize and cause ecological, economic and social damage. So far, no comprehensive analysis of the global accumulation and exchange of alien plant species between continents has been performed, primarily because of a lack of data. Here we bridge this knowledge gap by using a unique global database on the occurrences of naturalized alien plant species in 481 mainland and 362 island regions. In total, 13,168 plant species, corresponding to 3.9% of the extant global vascular flora, or approximately the size of the native European flora, have become naturalized somewhere on the globe as a result of human activity. North America has accumulated the largest number of naturalized species, whereas the Pacific Islands show the fastest increase in species numbers with respect to their land area. Continents in the Northern Hemisphere have been the major donors of naturalized alien species to all other continents. Our results quantify for the first time the extent of plant naturalizations worldwide, and illustrate the urgent need for globally integrated efforts to control, manage and understand the spread of alien species.

Remoteness promotes biological invasions on islands worldwide.

One of the best-known general patterns in island biogeography is the species-isolation relationship (SIR), a decrease in the number of native species with increasing island isolation that is linked to lower rates of natural dispersal and colonization on remote oceanic islands. However, during recent centuries, the anthropogenic introduction of alien species has increasingly gained importance and altered the composition and richness of island species pools. We analyzed a large dataset for alien and native plants, ants, reptiles, mammals, and birds on 257 (sub) tropical islands, and showed that, except for birds, the number of naturalized alien species increases with isolation for all taxa, a pattern that is opposite to the negative SIR of native species. We argue that the reversal of the SIR for alien species is driven by an increase in island invasibility due to reduced diversity and increased ecological naiveté of native biota on the more remote islands.

Global rise in emerging alien species results from increased accessibility of new source pools.

Our ability to predict the identity of future invasive alien species is largely based upon knowledge of prior invasion history. Emerging alien species-those never encountered as aliens before-therefore pose a significant challenge to biosecurity interventions worldwide. Understanding their temporal trends, origins, and the drivers of their spread is pivotal to improving prevention and risk assessment tools. Here, we use a database of 45,984 first records of 16,019 established alien species to investigate the temporal dynamics of occurrences of emerging alien species worldwide. Even after many centuries of invasions the rate of emergence of new alien species is still high: One-quarter of first records during 2000-2005 were of species that had not been previously recorded anywhere as alien, though with large variation across taxa. Model results show that the high proportion of emerging alien species cannot be solely explained by increases in well-known drivers such as the amount of imported commodities from historically important source regions. Instead, these dynamics reflect the incorporation of new regions into the pool of potential alien species, likely as a consequence of expanding trade networks and environmental change. This process compensates for the depletion of the historically important source species pool through successive invasions. We estimate that 1-16% of all species on Earth, depending on the taxonomic group, qualify as potential alien species. These results suggest that there remains a high proportion of emerging alien species we have yet to encounter, with future impacts that are difficult to predict.

Widespread decline in Central European plant diversity across six decades.

Based on plant occurrence data covering all parts of Germany, we investigated changes in the distribution of 2136 plant species between 1960 and 2017. We analyzed 29 million occurrence records over an area of ~350,000 km2 on a 5 × 5 km grid using temporal and spatiotemporal models and accounting for sampling bias. Since the 1960s, more than 70% of investigated plant species showed declines in nationwide occurrence. Archaeophytes (species introduced before 1492) most strongly declined but also native plant species experienced severe declines. In contrast, neophytes (species introduced after 1492) increased in their nationwide occurrence but not homogeneously throughout the country. Our analysis suggests that the strongest declines in native species already happened in the 1960s-1980s, a time frame in which often few data exist. Increases in neophytic species were strongest in the 1990s and 2010s. Overall, the increase in neophytes did not compensate for the loss of other species, resulting in a decrease in mean grid cell species richness of -1.9% per decade. The decline in plant biodiversity is a widespread phenomenon occurring in different habitats and geographic regions. It is likely that this decline has major repercussions on ecosystem functioning and overall biodiversity, potentially with cascading effects across trophic levels. The approach used in this study is transferable to other large-scale trend analyses using heterogeneous occurrence data.

Projecting the continental accumulation of alien species through to 2050.

Biological invasions have steadily increased over recent centuries. However, we still lack a clear expectation about future trends in alien species numbers. In particular, we do not know whether alien species will continue to accumulate in regional floras and faunas, or whether the pace of accumulation will decrease due to the depletion of native source pools. Here, we apply a new model to simulate future numbers of alien species based on estimated sizes of source pools and dynamics of historical invasions, assuming a continuation of processes in the future as observed in the past (a business-as-usual scenario). We first validated performance of different model versions by conducting a back-casting approach, therefore fitting the model to alien species numbers until 1950 and validating predictions on trends from 1950 to 2005. In a second step, we selected the best performing model that provided the most robust predictions to project trajectories of alien species numbers until 2050. Altogether, this resulted in 3,790 stochastic simulation runs for 38 taxon-continent combinations. We provide the first quantitative projections of future trajectories of alien species numbers for seven major taxonomic groups in eight continents, accounting for variation in sampling intensity and uncertainty in projections. Overall, established alien species numbers per continent were predicted to increase from 2005 to 2050 by 36%. Particularly, strong increases were projected for Europe in absolute (+2,543 ± 237 alien species) and relative terms, followed by Temperate Asia (+1,597 ± 197), Northern America (1,484 ± 74) and Southern America (1,391 ± 258). Among individual taxonomic groups, especially strong increases were projected for invertebrates globally. Declining (but still positive) rates were projected only for Australasia. Our projections provide a first baseline for the assessment of future developments of biological invasions, which will help to inform policies to contain the spread of alien species.

Drivers of future alien species impacts: An expert-based assessment.

Understanding the likely future impacts of biological invasions is crucial yet highly challenging given the multiple relevant environmental, socio-economic and societal contexts and drivers. In the absence of quantitative models, methods based on expert knowledge are the best option for assessing future invasion trajectories. Here, we present an expert assessment of the drivers of potential alien species impacts under contrasting scenarios and socioecological contexts through the mid-21st century. Based on responses from 36 experts in biological invasions, moderate (20%-30%) increases in invasions, compared to the current conditions, are expected to cause major impacts on biodiversity in most socioecological contexts. Three main drivers of biological invasions-transport, climate change and socio-economic change-were predicted to significantly affect future impacts of alien species on biodiversity even under a best-case scenario. Other drivers (e.g. human demography and migration in tropical and subtropical regions) were also of high importance in specific global contexts (e.g. for individual taxonomic groups or biomes). We show that some best-case scenarios can substantially reduce potential future impacts of biological invasions. However, rapid and comprehensive actions are necessary to use this potential and achieve the goals of the Post-2020 Framework of the Convention on Biological Diversity.

Similar factors underlie tree abundance in forests in native and alien ranges.

AIM: Alien plant species can cause severe ecological and economic problems, and therefore attract a lot of research interest in biogeography and related fields. To identify potential future invasive species, we need to better understand the mechanisms underlying the abundances of invasive tree species in their new ranges, and whether these mechanisms differ between their native and alien ranges. Here, we test two hypotheses: that greater relative abundance is promoted by (a) functional difference from locally co-occurring trees, and (b) higher values than locally co-occurring trees for traits linked to competitive ability. LOCATION: Global. TIME PERIOD: Recent. MAJOR TAXA STUDIED: Trees. METHODS: We combined three global plant databases: sPlot vegetation-plot database, TRY plant trait database and Global Naturalized Alien Flora (GloNAF) database. We used a hierarchical Bayesian linear regression model to assess the factors associated with variation in local abundance, and how these relationships vary between native and alien ranges and depend on species' traits. RESULTS: In both ranges, species reach highest abundance if they are functionally similar to co-occurring species, yet are taller and have higher seed mass and wood density than co-occurring species. MAIN CONCLUSIONS: Our results suggest that light limitation leads to strong environmental and biotic filtering, and that it is advantageous to be taller and have denser wood. The striking similarities in abundance between native and alien ranges imply that information from tree species' native ranges can be used to predict in which habitats introduced species may become dominant.

Naturalization of European plants on other continents: The role of donor habitats.

The success of European plant species as aliens worldwide is thought to reflect their association with human-disturbed environments. However, an explicit test including all human-made, seminatural and natural habitat types of Europe, and their contributions as donor habitats of naturalized species to the rest of the globe, has been missing. Here we combine two databases, the European Vegetation Checklist and the Global Naturalized Alien Flora, to assess how human influence in European habitats affects the probability of naturalization of their plant species on other continents. A total of 9,875 native European vascular plant species were assigned to 39 European habitat types; of these, 2,550 species have become naturalized somewhere in the world. Species that occur in both human-made habitats and seminatural or natural habitats in Europe have the highest probability of naturalization (64.7% and 64.5% of them have naturalized). Species associated only with human-made or seminatural habitats still have a significantly higher probability of becoming naturalized (41.7% and 28.6%, respectively) than species confined to natural habitats (19.4%). Species associated with arable land and human settlements were recorded as naturalized in the largest number of regions worldwide. Our findings highlight that plant species' association with native-range habitats disturbed by human activities, combined with broad habitat range, play an important role in shaping global patterns of plant invasions.

Conventional land-use intensification reduces species richness and increases production: A global meta-analysis.

Most current research on land-use intensification addresses its potential to either threaten biodiversity or to boost agricultural production. However, little is known about the simultaneous effects of intensification on biodiversity and yield. To determine the responses of species richness and yield to conventional intensification, we conducted a global meta-analysis synthesizing 115 studies which collected data for both variables at the same locations. We extracted 449 cases that cover a variety of areas used for agricultural (crops, fodder) and silvicultural (wood) production. We found that, across all production systems and species groups, conventional intensification is successful in increasing yield (grand mean + 20.3%), but it also results in a loss of species richness (-8.9%). However, analysis of sub-groups revealed inconsistent results. For example, small intensification steps within low intensity systems did not affect yield or species richness. Within high-intensity systems species losses were non-significant but yield gains were substantial (+15.2%). Conventional intensification within medium intensity systems revealed the highest yield increase (+84.9%) and showed the largest loss in species richness (-22.9%). Production systems differed in their magnitude of richness response, with insignificant changes in silvicultural systems and substantial losses in crop systems (-21.2%). In addition, this meta-analysis identifies a lack of studies that collect robust biodiversity (i.e. beyond species richness) and yield data at the same sites and that provide quantitative information on land-use intensity. Our findings suggest that, in many cases, conventional land-use intensification drives a trade-off between species richness and production. However, species richness losses were often not significantly different from zero, suggesting even conventional intensification can result in yield increases without coming at the expense of biodiversity loss. These results should guide future research to close existing research gaps and to understand the circumstances required to achieve such win-win or win-no-harm situations in conventional agriculture.

The role of fruit heteromorphism in the naturalization of Asteraceae.

BACKGROUND AND AIMS: Fruit heteromorphism is considered to be a bet-hedging strategy to cope with spatially or temporally heterogeneous environments. The different behaviours of the fruit morphs of the same species might also be beneficial during naturalization, once the species has been introduced to a new range. Yet, no study to date has tested the association between fruit heteromorphism and global-scale naturalization success for a large set of plant species. METHODS: We compiled two large datasets on fruit heteromorphism in Asteraceae. One dataset was on native species in Central Europe (n = 321) and the other was on species frequently planted as ornamentals (n = 584). Using phylogenetic linear and logistic regressions, we tested whether heteromorphic species are more likely to naturalize outside their native range, and in more regions of the world than monomorphic species. We also tested whether the effect of heteromorphism is modulated by life history and height of the species. KEY RESULTS: We show that heteromorphic species were more likely to naturalize outside their native range. However, among the naturalized species, heteromorphic and monomorphic species did not differ in the number of world regions where they became naturalized. A short life span and tall stature both promoted naturalization success and, when life history and height were included in the models, the effect of fruit heteromorphism on the ability to naturalize became non-significant. Nevertheless, among tall plants, heteromorphic ornamental species were significantly more likely to become naturalized in general and in more regions than monomorphic species. CONCLUSIONS: Our results provide evidence that in Asteraceae the production of heteromorphic fruits is associated with naturalization success. It appears, however, that not fruit heteromorphism per se, but a successful combination of other biological traits in fruit heteromorphic species, namely short life span and tall stature, contributes to their naturalization success.

Global mismatches in aboveground and belowground biodiversity.

Human activities are accelerating global biodiversity change and have resulted in severely threatened ecosystem services. A large proportion of terrestrial biodiversity is harbored by soil, but soil biodiversity has been omitted from many global biodiversity assessments and conservation actions, and understanding of global patterns of soil biodiversity remains limited. In particular, the extent to which hotspots and coldspots of aboveground and soil biodiversity overlap is not clear. We examined global patterns of these overlaps by mapping indices of aboveground (mammals, birds, amphibians, vascular plants) and soil (bacteria, fungi, macrofauna) biodiversity that we created using previously published data on species richness. Areas of mismatch between aboveground and soil biodiversity covered 27% of Earth's terrestrial surface. The temperate broadleaf and mixed forests biome had the highest proportion of grid cells with high aboveground biodiversity but low soil biodiversity, whereas the boreal and tundra biomes had intermediate soil biodiversity but low aboveground biodiversity. While more data on soil biodiversity are needed, both to cover geographic gaps and to include additional taxa, our results suggest that protecting aboveground biodiversity may not sufficiently reduce threats to soil biodiversity. Given the functional importance of soil biodiversity and the role of soils in human well-being, soil biodiversity should be considered further in policy agendas and conservation actions by adapting management practices to sustain soil biodiversity and considering soil biodiversity when designing protected areas.

Disparidades Mundiales entre la Biodiversidad Sobre y Bajo el Suelo Resumen Las actividades humanas están acelerando el cambio en la biodiversidad mundial y han tenido como resultado unos servicios ambientales severamente amenazados. Una gran proporción de la biodiversidad terrestre está albergada en el suelo, pero la biodiversidad de este ha sido omitida de varias evaluaciones mundiales de biodiversidad y de las acciones de conservación, además de que el entendimiento de los patrones mundiales de la biodiversidad del suelo permanece limitado; particularmente, la extensión del traslape entre los puntos fríos y calientes de biodiversidad sobre y bajo suelo no está clara. Examinamos los patrones mundiales de estos traslapes mapeando los índices de biodiversidad sobre el suelo (mamíferos, aves, anfibios y plantas vasculares) y bajo el suelo (bacterias, hongos y macrofauna) que creamos con datos previamente publicados de la riqueza de especies. Las áreas de disparidad entre la biodiversidad sobre y bajo el suelo cubrieron el 27% de la superficie terrestre del planeta. El bioma de los bosques templados de plantas frondosas y mixtas tuvo la proporción más alta de celdas de cuadrícula con una biodiversidad alta sobre el suelo, pero baja para en el subsuelo, mientras que los biomas boreales y de la tundra tuvieron una biodiversidad intermedia bajo el suelo, pero baja para el sobre suelo. Aunque se requieren más datos sobre la biodiversidad del suelo, tanto para cubrir los vacíos geográficos como para incluir a taxones adiciones, nuestros resultados sugieren que la protección a la biodiversidad sobre el suelo puede no reducir suficientemente las amenazas para la biodiversidad del suelo. Dada la importancia funcional de la biodiversidad del suelo y el papel de los suelos en el bienestar humano, se debería considerar a la biodiversidad del suelo mucho más en las agendas políticas y en las acciones de conservación, adaptando a las prácticas de manejo para que mantengan a la biodiversidad del suelo y la consideren cuando designen áreas protegidas.

The role of adaptive strategies in plant naturalization.

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