A peer-reviewed open-access journal NeoBiota 91: 29-48 (2024) doi: 10.3897/neobiota.91.1 15675 RESEARCH ARTICLE &) NeoBiota https:/ / neobi ota. pen soft. net Advancing research on alien species and biological invasions Clonal alien plants in the mountains spread upward more extensively and faster than non-clonal Miao-Miao Zheng!”, Petr PySek**, Kun Guo!”, Hasigerili'’, Wen-Yong Guo'?° | Zhejiang Tiantong Forest Ecosystem National Observation and Research Station, School of Ecological and Environmental Sciences, East China Normal University, Shanghai, 200241, China 2. Research Center for Global Change and Complex Ecosystems, School of Ecological and Environmental Sciences, East China Normal University, Shanghai, 200241, China 3 Czech Academy of Sciences, Institute of Botany, Department of Invasion Ecology, Prihonice, CZ-25243, Czech Republic 4 Department of Ecology, Faculty of Science, Charles University, Vinitnd 7, Prague, CZ-12844, Czech Republic 5 Shanghai Key Lab for Urban Ecological Processes and Eco-Restoration, School of Ecological and Environmental Sciences, East China Normal University, Shanghai, 200241, China Corresponding author: Wen-Yong Guo (wyguo@des.ecnu.edu.cn) Academic editor: Gerhard Karrer | Received 13 November 2023 | Accepted 22 January 2024 | Published 8 February 2024 Citation: Zheng M-M, PySek PB, Guo K, Hasigerili, Guo W-Y (2024) Clonal alien plants in the mountains spread upward more extensively and faster than non-clonal. NeoBiota 91: 29-48. https://doi.org/10.3897/neobiota.91.115675 Abstract Alien species are colonizing mountain ecosystems and increasing their elevation ranges in response to ongoing climate change and anthropogenic disturbances, posing increasing threats to native species. How- ever, how quickly alien species spread upward and what drives their invasion remains insufficiently under- stood. Here, using 26,952 occurrence records of 58 alien plant species collected over two centuries in the Czech Republic, we explored the elevation range and invasion speed of each alien species and the underly- ing factors driving these variables. We collected species traits relevant for invasion (e.g., clonality, flower- ing time, life span, invasion status, height, mycorrhizal type, native range, naturalized range, monoploid genome size, and Ellenberg-type indicator values for light, temperature, and nitrogen), human-associated factors (e.g., introduction pathways and the sum of economic use types), and minimum residence time. We explored the relationships between these factors and species’ elevation range and invasion speed using phylogenetic regressions. Our results showed that 58 alien species have been expanding upward along mountain elevations in the Czech Republic over the past two centuries. A stronger effect of species’ traits than human-associated factors has been revealed, e.g., clonality was a key trait supporting the invasion of alien species into the mountains, while human-associated factors showed no effect. Our findings highlight that the characteristics associated with rapid reproduction and spread are crucial for alien species’ invasion into montane regions. Identifying key drivers of this process is important for predicting the spatiotempo- ral dynamics of alien species in high-altitude ecosystems and thus employing apposite measures to reduce the threat to native plant species. Copyright Miao-Miao Zheng et al. This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited. 30 Miao-Miao Zheng et al. / NeoBiota 91: 29-48 (2024) Keywords Alien plant species, clonal trait, elevation range, human activities, invasion speed, mountain invasions Introduction Mountains are of crucial importance for biodiversity conservation (Rangel et al. 2018; Rahbek et al. 2019). Conventionally, these regions were considered to be relatively unharmed by alien species (McDougall et al. 2011; Alexander et al. 2016). However, with intensifying climate change and human pressure, alien species are now colonizing mountains and increasing their elevation range upward, posing a significant threat to native species (Pauchard et al. 2009; McDougall et al. 2011; PySek et al. 2011; Alexan- der et al. 2016; Dainese et al. 2017; Koide et al. 2017). Consequently, a comprehensive understanding of the increasing trends in alien species distribution and the speed at which alien species are establishing at different elevations becomes imperative for effec- tive conservation strategies in mountain regions (Iseli et al. 2023). In mountainous areas, the influx of alien species typically commences at lower elevations before gradually spreading upward over time (Alexander et al. 2011; PySek et al. 2011; Marini et al. 2013). The directional ecological filtering hypothesis, a phenomenon of a decline in alien species richness with increasing elevation due to a progressive species loss, is often invoked to elucidate the distribution of alien flora along altitudinal gradients worldwide (Alexander et al. 2011). In addition, the up- ward expansion of alien species is associated with minimum residence time (PySek et al. 2011). Recent investigations have underscored its significant role in shaping the potential elevation range of alien species (Pysek et al. 2009a, 2011, 2015; Alex- ander et al. 2011), because longer residence time allows for more extensive dispersal or can result in genetic adaptation (Becker et al. 2005; Haider et al. 2010; PySek et al. 2011). In addition to the minimum residence time, various characteristics of alien spe- cies, e.g., introduction pathways (Alexander et al. 2011), economic utility (Balestri et al. 2018, van Kleunen et al. 2020), proximity to road networks (Dainese et al. 2017; Skalova et al. 2017), as well as their inherent traits (e.g., dispersal abilities, climatic adaptability, and genetic adaptation) (Dietz and Edwards 2006; Alexander 2010) have been recognized as key drivers of species’ range expansion. Theoretically, deliberately introduced and economically valuable plants (Pergl et al. 2017; Balestri et al. 2018; van Kleunen et al. 2018) often exhibit larger propagule pressure in terms of both propagule quantity and frequency of introduction events, consequently enhancing the likelihood of successful establishment. In addition, species with rapid growth, short generations, and strong competitive ability are also more likely to become successful invaders (van Kleunen et al. 2010). For instance, plant height was found to be positively related to species’ range and probability of invasion (PySek et al. 2009a, 2015; Divisek et al. 2018); optimal flowering time, functioning as a reproductive trait, ensures seed fertility Clonal species invade larger and faster og (Celesti-Grapow et al. 2003; Godoy et al. 2009), and species with earlier flowering can avoid competition and expedite the life cycle, leading to prompt reproduction. On a macroecological scale, clonality was reported to play a positive role in the invasion suc- cess and distribution of alien species (Pysek 1997; Liu et al. 2006; Wang et al. 2024) and species primarily engaging in asexual reproduction also tend to exhibit broader ranges (Cosendai et al. 2013). Moreover, studies have shed light on how the geographic expansion of alien plants benefits from the presence of mycorrhizal associations, which form specialized structures aiding vegetative dispersal (Menzel et al. 2017; Correia et al. 2018; PySek et al. 2019). Karyological characteristics were recently suggested as an important trait under- pinning plant invasion success, and species with small genomes proved to be at an advantage in the process of alien plants’ naturalization (KubeSova et al. 2010; Suda et al. 2015; Lopes et al. 2021; PySek et al. 2023) as the “large genome constraints” proposed (Knight et al. 2005). However, the advantage of possessing a small genome does not translate into the more advanced stage of the invasion process characterized by the rapid spread; here, the opposite is true as species with relatively large genomes are more likely to be successful invaders (Lopes et al. 2021; Carta et al. 2022; PySek et al. 2023). While the magnitude of a species’ native range significantly influences its pread- aptation to the introduced environment, and a broader native range generally fosters greater readiness for the establishment and thriving within the novel environment (PySek et al. 2015; Guo et al. 2019; Fristoe et al. 2023), the inclusion of species-specific environmental preferences, e.g., optimal values or ranges for existence, development, growth, and reproduction, can offer a more nuanced understanding of species’ spatial expansion patterns, particularly along environmental gradients like elevation (Di Biase et al. 2023). Despite these insights, scant knowledge exists regarding the key driving forces behind changes in the elevation range or vertical spread of alien plants (Dainese et al. 2017; Auld et al. 2022), especially when considering the relatively low propagule pressure at high elevations (Alexander et al. 2011). In this study, we aim to estimate the speed at which alien species increase their elevation range and identify the drivers that underlie such invasions. Specifically, we calculated the elevation range and invasion speed of 58 alien species for which such data exist in the regional dataset for the Czech Republic. We also used various char- acteristics of these plants, including both species’ inherent traits (e.g., clonality, flow- ering time, monoploid genome size, and Ellenberg-type indicator values reflecting ecological demands) and human-associated factors (e.g., introduction pathways and economic use types). We then used phylogenetic regressions to identify the key driv- ers of elevation range and invasion speed. We hypothesized that: 1) alien species are expanding upward along mountain elevations in the Czech Republic; 2) both species’ inherent traits and human-associated factors contribute to the upward of alien spe- cies, and play distinct roles in determining the elevational range (and changes) and invasion speed. 32 Miao-Miao Zheng et al. / NeoBiota 91: 29-48 (2024) Materials and methods Species distribution records Species data were obtained from an existing dataset (Williamson et al. 2005; Pysek et al. 2011), originally including 65 alien species introduced to the Czech Republic after 1500 AD. The dataset compiled the elevation of alien species in the mountains since the year 1738, with 28,288 occurrence records in total. Based on this data, we calcu- lated each species’ elevation range (Elevation , — Elevation ,,.) (m), invasion speed ((Elevation | — Elevation )/ (year )) (m/year), and the minimum residence time (2022 — year — year year_of_1st_record max_elevation 1st_record lst_record? * From the complete dataset, we removed abnormal values of invasion speed, e.g., values for species with no elevation record in the year of first introduction; values for species for which there was less than 40 years since their first introduction to the maxi- mum elevation year, due to the possibility of inadequate sampling; and negative values. We used the ‘WorldFlora R package (Kindt 2020) to standardize all taxon names be- fore matching species trait data further. Species traits We obtained data on the clonality (modular species with potential vegetative reproduc- tion or unitary species without this potential), flowering time (as the first month of flowering), life span (annual, perennial, or both), and invasion status (casual, natural- ized, invasive) of the 58 alien species from existing datasets (Williamson et al. 2005; Pysek et al. 2022). The categories of invasion status are based on a well-defined frame- work (Blackburn et al. 2011): alien species that do not form self-sustaining popula- tions in the introduced regions are casuals; alien species that form self-sustaining popu- lations for several life cycles in the introduced region without human intervention are naturalized; and alien species that can maintain self-replacing populations at consider- able distances from their parents and/or sites of introduction, producing reproductive offspring and having the potential to spread over long distances are invasive. The height of each species was extracted from the LEDA database (https://uol. de/en/landeco/research/leda) (Kleyer et al. 2008) using an average method. We then compiled Ellenberg-type indicator values for light, temperature, and nitrogen for each species, which are expert-based rankings of plant species according to their ecological optima on main environmental gradients (recently updated and harmonized: Tichy et al. 2022). Smaller Ellenberg-type indicator values represent species that are better adapted to the lower levels of a given factor. Mycorrhizal types were obtained from the FungalRoot v.2.0 database (Soudzilovskaia et al. 2022), divided into arbuscular mycor- rhizal (AM) and non-mycorrhizal (NM) plant species. Native range data for alien species were extracted from the POWO (Plants of the World Online; https://powo.science.kew.org/) database, and naturalized range Clonal species invade larger and faster 33 data were obtained from the GloNAF (Global Naturalized Alien Flora) database (van Kleunen et al. 2015, 2019; Pysek et al. 2017). These two range-related data were ex- pressed as the number of the Taxonomic Database Working Group (TDWG) level 3 regions (level 3 corresponds to “Botanical Countries”) in which species are recorded as native or naturalized, respectively (Brummitt, 2001). Monoploid genome sizes (i.e., DNA content in a single chromosome set, sensu Greilhuber et al. (2005)) for the spe- cies were filtered from the Plant DNA C-values (https://cvalues.science.kew.org/) (Pel- licer and Leitch 2020). Human-associated factors Information on the introduction pathway (deliberate or accidental introduction) of each alien species to the Czech Republic was taken from PySek et al. (2012). Then, we collated the economic use data of each species from the WCUP (World Checklist of Useful Plant Species; Diazgranados et al. 2020). The WCUP list provides informa- tion on 10 distinct economic uses, such as medicines and materials. We calculated the overall sum of economic use types for each species and set the economic use of species not included in the list as zero. Phylogenetic tree To consider potential phylogenetic relatedness across the species, a phylogenetic tree was created for the 58 species using the ‘V-PhyloMaker’ package based on the default setting (Jin and Qian 2019). This phylogenetic tree was used to impute the missing traits values and subsequent statistical analyses. Imputation of missing trait values Given the presence of missing data for certain species, we undertook gap-filling pro- cesses as follows. For the 20 species lacking monoploid genome size, imputation was carried out utilizing the full dataset of 12,273 species (Pellicer and Leitch 2020) via the R package ‘“Rphylopars (Goolsby et al. 2017). The 11 species lacking Ellenberg-type indicator values and four species without mycorrhizal types were supplemented by data from the same genus, given the strong phylogenetic conservatism for these indicators (Prinzing et al. 2001). Finally, a list of 58 alien plant species with 26,952 occurrence records and complete values was obtained, and 45 species for the invasion speed. Statistical analyses All data analyses and visualizations were performed in R v4.2.1 (R Core Team 2023). First, Pearson correlation analysis was performed with all the continuous variables (i.e., invasion speed, flowering time, plant height, Ellenberg indicator values for light, 34 Miao-Miao Zheng et al. / NeoBiota 91: 29-48 (2024) temperature and nutrients, native range size, naturalized range size, the sum of eco- nomic uses, monoploid genome size, and minimum residence time), which showed no strong collinearity among the variables (Appendix 1: Fig. Al; r < 0.7). Plant height, native range size, naturalized range size, and monoploid genome size were log-trans- formed to approximate normal distribution. To examine whether the elevation range and invasion speed varied across the categories of clonality, invasion status, life span, mycorrhizal types, or introduction pathways, phylogenetic ANOVA models were used with function ‘phy/ANOVA in R package ‘phytools (Garland et al. 1993; Revell 2012). Phylogeny is considered in the analysis because species may not represent statistically independent data points, thus avoiding unreliable model estimates (Garland et al. 1993). P-values were determined by Brownian motion model simulation phylogeny, which was run 10,000 times. To explore the combined effects of continuous variables on elevation range and invasion speed of alien species, phylogenetic multiple linear re- gression models were employed (Si et al. 2022). Models were run separately for clonal and non-clonal species and the variables selection was conducted using the ‘phylostep’ function in the ‘“Phylolm’ package (Tung Ho and Ané 2014). All numerical variables were standardized to gain standardized coefficients so that comparisons between and within models could be possible (Schielzeth 2010). Finally, according to the results of the ‘phylostep’, several predictors were identified as key drivers for both clonal and non-clonal alien species, which were used in regressions on data of clonal species, non- clonal species, and all species. Results The elevation ranges of the 58 alien species varied from 132 m to 1095 m (Fig. 1). Heracleum mantegazzianum had the largest elevation range (1095 m) and the highest upper elevation limit, peaking at 1294 m a.s.l. In contrast, Panicum dichotomiflorum had the smallest elevation range (132 m) and the lowest upper elevation limit with a peak of 271 m a.s.l. As for the elevational invasion speed of the 45 alien plants ana- lyzed, it was fastest for Mimulus guttatus with 8.4 m/year, whereas for Ambrosia arte- misiifolia it was the slowest (0.5 m/year) (Fig. 1). Generally, species with large elevation ranges also had high invasion speeds (Appendix 1: Fig. A1). Both elevation range and invasion speed differed with regard to clonality (Fig. 2a, c; P< 0.01). Specifically, clonal species had significantly larger elevation ranges (aver- age values of clonal vs. non-clonal species = 709 m vs. 500 m, P = 0.001) and higher invasion speeds (5.4 vs. 3.7 m/year, P = 0.004) than non-clonal species (Fig. 2a, c). Casual species had the smallest elevation ranges compared to both naturalized and invasive species (357 m, 613 m, and 688 m on average, respectively, P < 0.05), which were not significantly different from each other (Fig. 2b; P > 0.05). Likewise, no significant differences were observed in invasion speed among the three categories of invasion status (Fig. 2d). Moreover, neither elevation range nor invasion speed Clonal species invade larger and faster 35 Ambrosia trifida Ambrosia artemisiifolia Xanthium spinosum Helianthus annuus Galinsoga parviflora Galinsoga ciliata Bidens frondosa Telekia speciosa Solidago gigantea Solidago canadensis Erigeron canadensis Artemisia verlotiorum Artemisia annua Lactuca tatarica Heracleum mantegazzianum Veronica filiformis Veronica persica Digitalis purpurea Erythranthe guttata Cuscuta campestris Impatiens parviflora Impatiens glandulifera Amaranthus powellii Amaranthus retroflexus Amaranthus blitoides Amaranthus viridis Amaranthus albus Dysphania pumilio Dysphania botrys Chenopodium foliosum Kochia scoparia Mirabilis nyctaginea Reynoutria sachalinensis Reynoutria japonica Rumex patientia Rumex triangulivalvis Hirschfeldia incana Sisymbrium volgense Bunias orientalis A butilon theophrasti Epilobium ciliatum Oenothera biennis Geranium pyrenaicum Echinocystis lobata Cannabis ruderalis Lupinus polyphyllus Trifolium hybridum Corydalis lutea Consolida orientalis Panicum miliaceum Panicum dichotomiflorum Panicum capillare Sorghum halepense Hordeum jubatum Juncus tenuis Commelina communis Sisyrinchium angustifolium Acorus calamus Eee ee 0 50 100 150 0) 500 1000 Phylogeny (million years) Elevation range (m) Invasion speed (m/year) Figure |. Phylogenetic tree of the examined 58 alien plant species in the Czech Republic, with their elevation ranges (m) and invasion speeds (available for 45 species, m/year) aligned. differed among different life spans, mycorrhizal types, and introduction pathways (Appendix 1: Fig. A2). Phylogenetic regressions revealed different factors driving the elevation range and invasion speed for clonal and non-clonal species (Figs 3, 4). Specifically, for both clonal and non-clonal species, species’ invasion speed and minimum residence time showed positive effects on elevation range, whereas Ellenberg indicator values for temperature and the time of flowering showed negative effects, with species demanding a higher temperature and those that start flowering early having small elevation ranges (Figs 3a, 4). As for elevational invasion speed, native range size was the only predictor showing a negative effect on the invasion speed of non-clonal species (Fig. 3b). For clonal spe- cies, the invasion speed was negatively associated with Ellenberg indicator values for temperature, and with minimum residence time, and positively to naturalized range size (Fig. 3b). 36 Miao-Miao Zheng et al. / NeoBiota 91: 29-48 (2024) P=0.001 P=0.01 (a) (b) [__] _P=0.003 1000 - = 900- 7 = t D c Lmean = 709.00 750 = wo La ‘I * 600- | = iL 5 AD Drrcan = 613.00 rs 500 - I ) 7 = r end Fr Dmean = 356.85 250 - Fig no yes casual naturalized invasive (n = 34) (n = 24) (n = 13) (n = 21) (n = 24) (c) P= 0.004 (d) _= 9 8 © © onal = 3- Toy © 7.5- ® = E 6- 3 f 2 5.0- Uimean = 4-51 n c aoe 2 n $ & 2.5- 7 no yes casual naturalized invasive (n = 27) (n = 18) (n= 5) (n = 18) (n = 22) Clonal Status Figure 2. Violin plots showing the elevation ranges and invasion speeds of the examined alien species, categorized based on their clonality and invasion status. Note that the invasion speed was only available for 45 species. For the boxplots inside each violin plot, the horizontal line, red dot, and box, respectively, represent the median, the mean, and the interquartile range. P-values were obtained from phylogenetic ANOVAs and showed in the upper part of the violin plot if significant (P < 0.05). Discussion Using more than 26,000 historical occurrence records of 58 alien plant species intro- duced to the Czech Republic after 1738, we found a substantial upward shift along the elevations for the majority of the species analyzed (Fig. 1). The speed of spread to the higher elevation ranged from 0.5 to 8.4 m/year, with species capable of reaching the highest elevation also exhibiting the fastest invasion speed (Fig. 1). Our findings are in line with a recent study showing that alien plant species were expanding their upper elevation limits in 10 out of the 11 surveyed mountains across five continents (Iseli et al. 2023). However, no upward expansion of alien plants was observed in several regional studies, e.g., 67% of naturalized invasive plant species in California showed no mean elevation shift over the past century (Wolf et al. 2016). On the island of Ha- waii, both the upper and lower elevation limit of 20 alien species are moving up, but elevation ranges did not change significantly over 40 years (Koide et al. 2017). In Eu- rope, 10% of alien species exhibited a potential downslope shift (Dainese et al. 2017). Although these contrasting findings may be due to different climates and land-use Clonal species invade larger and faster 37 Elevation range (m) Invasion speed (m/year) Flowering time Ellenberg indicator values of temperature Native range Minimum residence time oo Height nf Naturalized range == ~~ Economic use sum Invasion speed Group Monoploi nome siz onoploid genome size =® non-clonal Ellenberg indicator =® clonal values of nutrients -1.0 -0.5 0.0 0.5 1.0 -1.0 -0.5 0.0 0.5 1.0 Estimate Figure 3. Standardized estimates and associated 95% confidence intervals obtained from phylogenetic re- gressions for each of the elevation range and invasion speed models. Models were run separately for clonal (orange) and non-clonal (blue) species and the variables were selected via the 'phylostep' function in the R package ‘Phylolm’. In particular, we added invasion speed as an additional variable for the elevation range. Confidence intervals that do not cross the zero line indicate that the estimates are significant (P < 0.05). history between the studying areas (Iseli et al. 2023), an understanding of the driving factors underlying the range expansions along elevation is emerging. Among the various factors tested in this study, alien species with clonal reproduc- tion exhibited not only significantly broader elevation ranges but also faster upward expansion compared to their non-clonal counterparts (Fig. 2a, c). Such a result is in line with previous studies showing that species primarily reproducing through asexual means tend to have broader distribution ranges (Cosendai et al. 2013). Besides, alien clonal species were found to have better growth performance than native clonal species, highlighting the pivotal role of clonality in promoting plant invasion (PySek 1997; Liu et al. 2019; Wang et al. 2019). Here, we also found that among clonal alien species, the taller ones exhibited a tendency for broader elevation ranges, and those with larger naturalized ranges had greater elevation ranges and speeds (Fig. 3a, b). For other traits tested with regard to the elevation expansion of alien species, we observed a negative association between monoploid genome size and eleva- tion range for clonal aliens (Fig. 3a). This can be explained by the “large genome constraints” hypothesis, stating that large genome acts as a constraint in extreme 38 Miao-Miao Zheng et al. / NeoBiota 91: 29-48 (2024) LI = -1 0 1 Ellenberg indicator values of temperature oO = ol Elevation range (m) eooae @ gt wo a a a Group =-®- non-clonal ) e =® clonal 8 e e R=0.42, p=0.0011 R=0.46, p = 0.00024 Dari Fer cach aL) eerie Flowering time Minimum residence time Figure 4. Relationships between elevation ranges and important variables identified from phylogenetic regressions (See Fig. 3 for details; data were log-transformed). Blue, orange, and black lines represent regression lines with non-clonal, clonal, and all species, respectively. The dashed lines represent non- significant relations. Grey shading represents 95% confidence intervals. and inhospitable conditions (Knight et al. 2005), which can be applied to alien species’ establishment in higher elevation. The invasion speed was strongly associ- ated with clonality, and for non-clonal alien species, only their native range size was negatively related to their upward spread (Fig. 3b). Although native range size was closely related to the fitness of alien species in the introduced areas (PySek et al. 2015; Guo et al. 2019; Fristoe et al. 2023), the inherent characteristics and interactions with local native species may be more important in determining the invasion speed of alien species. In essence, these results highlight the multifaceted interplay between clonality, height, genetic characteristics, and climate niche, re- vealing the intricate dynamics that contribute to the upward expansion and inva- siveness of clonal alien plants. In addition, our study identified the minimum residence time, flowering time, and demands for temperature as key factors driving the elevation range of alien species in the Czech Republic, regardless of their clonality status (Fig. 3a). Such results indicate that alien plants which are more likely to have a wider elevation range are those that Clonal species invade larger and faster 39 start flowering earlier, arrive earlier and have a reduced dependence on temperature conditions. These characteristics are also commonly observed in alpine plants (Alex- ander et al. 2016; Inouye 2020; Wang et al. 2020) and contribute to the expansion of their ecological range. It is gradually recognized that the position on the introduction-naturalization- invasion continuum is a good indicator of species’ minimum residence time (Pysek et al. 2009b; Moodley et al. 2013), thus aligning with the minimum residence time results: the elevation ranges of the 58 alien species increased along the introduction- naturalization-invasion continuum (Fig. 2b). While the same trend along the invasion continuum was not obtained for the invasion speed of alien species, a negative relation- ship was uncovered between the minimum residence time and the invasion speed for clonal species. The suitable ecological niche of the species may be responsible for this phenomenon, i.e., alien species with longer minimum residence time may have already occupied the most suitable habitats and, therefore, have slower invasion speeds. On the other hand, the relatively limited availability of valid invasion speed data for the initial two stages, in comparison to the third stage, might have introduced a degree of compromise to the statistical analysis. This potential data limitation could lead to the nuanced results observed in this context. For human-associated factors, we hypothesized that the economic use of alien plants will positively correlate with their elevation ranges, in keeping with the well- established assertions of earlier research, which underline the significance of economic use in determining the success of alien plant species within introduced ranges (Guo et al. 2019, van Kleunen et al. 2020). However, our results did not match the second hypothesis we proposed. ‘This may be due to nearly a quarter of alien species not having economic use data. Similarly, the results for the introduction pathway had no signifi- cant effect on driving alien species expansion along elevation (Appendix 1: Fig. A2). This could be caused by other human-related factors which could have greater predict- ability, such as distance to roads. Proximity to roads proved to be a key driver of the observed rapid upward spread of alien species (Dainese et al. 2017). Although we considered several factors related to species’ expansion along the el- evation gradient, several important variables, such as climate and soil properties, were missing from the analysis. It appears necessary to include these variables in future stud- ies to gain a more comprehensive understanding. Interactions between alien and native species are equally important to become a subject of future studies, with the potential to provide valuable insights into the mechanisms underlying the establishment and persistence of alien species in alpine habitats. In summary, our results showed that 58 alien species have been expanding upward along mountain elevations in the Czech Republic over the past two centuries. Alien species can reach the highest elevations and exhibit the widest range of elevations, providing further support for the hypothesis of directional ecological filtering. In par- ticular, our study explored how species traits and human-associated factors influence the elevation range and invasion speed of alien species towards mountains. We found distinct roles of species characteristics and human-associated factors in shaping species’ 40 Miao-Miao Zheng et al. / NeoBiota 91: 29-48 (2024) elevational expansion, e.g., compared with non-clonal alien species, clonal alien species had a wider elevation range and faster invasion speed, while human-associated factors had no effect. Our results emphasized that rapid reproduction and spread are crucial for alien species’ expansion in mountainous regions and are further facilitated by long residence time. Identifying key drivers of the distribution and spread of alien species in mountain areas and further developing a more complete understanding of how traits, human factors, and climate interact is critical. By analyzing complex temporal patterns and trends in the distribution of alien species, we can better grasp their dynamics and potential impacts on local ecosystems given the dynamic climate change worldwide. Acknowledgments MMZ, KG, H, and WYG were supported by the Shanghai Pujiang Program (grant 21PJ1402700 awarded to WYG). PP was supported by EXPRO grant no. 19-28807X (Czech Science Foundation) and long-term research development project RVO 67985939 (Czech Academy of Sciences). KG was supported by the Shanghai Sailing Program (grant 22YF1411700). References Alexander JM (2010) Genetic differences in the elevational limits of native and introduced Lactuca serriola populations. Journal of Biogeography 37(10): 1951-1961. https://doi. org/10.1111/j.1365-2699.2010.02335.x Alexander JM, Kueffer C, Daehler CC, Edwards PJ, Pauchard A, Seipel T, Arévalo J, Cavieres L, Dietz H, Jakobs G, McDougall K, Naylor B, Otto R, Parks CG, Rew L, Walsh N (2011) Assembly of nonnative floras along elevational gradients explained by directional ecological filtering. Proceedings of the National Academy of Sciences of the United States of America 108(2): 656-661. https://doi.org/10.1073/pnas. 1013136108 Alexander JM, Lembrechts JJ, Cavieres LA, Daehler C, Haider S, Kueffer C, Liu G, McDougall K, Milbau A, Pauchard A, Rew LJ, Seipel T (2016) Plant invasions into mountains and alpine ecosystems: Current status and future challenges. Alpine Botany 126(2): 89-103. https://doi.org/10.1007/s00035-016-0172-8 Auld J, Everingham SE, Hemmings FA, Moles AT (2022) Alpine plants are on the move: Quantifying distribution shifts of Australian alpine plants through time. Diversity & Distributions 28(5): 943-955. https://doi.org/10.1111/ddi.13494 Balestri E, Vallerini F, Menicagli V, Barnaba S, Lardicci C (2018) Biotic resistance and vegeta- tive propagule pressure co-regulate the invasion success of a marine clonal macrophyte. Scientific Reports 8(1): 16621. https://doi.org/10.1038/s41598-018-35015-0 Becker T, Dietz H, Billeter R, Buschmann H, Edwards PJ (2005) Altitudinal distribution of al- ien plant species in the Swiss Alps. Perspectives in Plant Ecology, Evolution and Systematics 7(3): 173-183. https://doi.org/10.1016/j.ppees.2005.09.006 Clonal species invade larger and faster 4] Blackburn TM, PySek P, Bacher S, Carlton JT, Duncan RP, Jarosik V, Wilson JRU, Richardson DM (2011) A proposed unified framework for biological invasions. ‘Trends in Ecology & Evolution 26(7): 333-339. https://doi.org/10.1016/j.tree.2011.03.023 Brummitt RK (2001) World geographical scheme for recording plant distributions. ed. 2. Hunt Institute for Botanical Documentation & Carnegie Mellon University, Pittsburgh, 137 pp. https://grassworld.myspecies.info/sites/grassworld.myspecies.info/files/tdwg_geo2.pdf Carta A, Mattana E, Dickie J, Vandelook F (2022) Correlated evolution of seed mass and ge- nome size varies among life forms in flowering plants. Seed Science Research 32(1): 46-52. https://doi.org/10.1017/S096025852200007 1 Correia M, Heleno R, Vargas P, Rodriguez-Echeverria S (2018) Should I stay or should I go? Mycorrhizal plants are more likely to invest in long-distance seed dispersal than non-myc- orrhizal plants. Ecology Letters 21(5): 683-691. https://doi.org/10.1111/ele.12936 Cosendai AC, Wagner J, Ladinig U, Rosche C, Horandl E (2013) Geographical parthenogen- esis and population genetic structure in the alpine species Ranunculus kuepferi (Ranuncu- laceae). Heredity 110(6): 560-569. https://doi.org/10.1038/hdy.2013.1 Dainese M, Aikio S$, Hulme PE, Bertolli A, Prosser F Marini L (2017) Human disturbance and upward expansion of plants in a warming climate. Nature Climate Change 7(8): 577-580. https://doi.org/10.1038/nclimate3337 Di Biase L, Tsafack N, Pace L, Fattorini S (2023) Ellenberg indicator values disclose com- plex environmental filtering processes in plant communities along an elevational gradient. Biology (Basel) 12(2): 161. https://doi.org/10.3390/biology12020161 Diazgranados M, Allkin B, Black N, Camara-Leret R, Canteiro C, Carretero J, Eastwood R, Hargreaves S, Hudson A, Milliken W, Nesbitt M, Ondo I, Patmore K, Pironon S, Turn- er R, Ulian T (2020) World Checklist of Useful Plant Species. https://doi.org/10.5063/ F1CV4G34 Dietz H, Edwards PJ (2006) Recognition that causal processes change during plant invasion helps explain conflicts in evidence. Ecology 87(6): 1359-1367. https://doi.org/10.1890/0012- 9658(2006)87[1359:RTCPCD]2.0.CO;2 Divisek J, Chytry M, Beckage B, Gotelli NJ, Lososova Z, PySek P, Richardson DM, Molofsky J (2018) Similarity of introduced plant species to native ones facilitates naturalization, but differences enhance invasion success. Nature Communications 9(1): 1-10. https://doi. org/10.1038/s41467-018-06995-4 Fristoe TS, Bleilevens J, Kinlock NL, Yang Q, Zhang Z, Dawson W, Essl F, Kreft H, Pergl J, Pysek P, Weigelt P, Dufour-Dror J-M, Sennikov AN, Wasowicz P, Westergaard KB, van Kleunen M (2023) Evolutionary imbalance, climate and human history jointly shape the global biogeography of alien plants. Nature Ecology & Evolution 7(10): 1633-1644. https://doi.org/10.1038/s41559-023-02172-z Garland T, Dickerman AW, Janis CM, Jones JA (1993) Phylogenetic analysis of covariance by computer simulation. Systematic Biology 42(3): 265-292. https://doi.org/10.1093/sys- bio/42.3.265 Godoy O, Richardson DM, Valladares F, Castro-Diez P (2009) Flowering phenology of invasive alien plant species compared with native species in three Mediterranean-type ecosystems. Annals of Botany 103(3): 485-494. https://doi.org/10.1093/aob/mcn232 42 Miao-Miao Zheng et al. / NeoBiota 91: 29-48 (2024) Goolsby EW, Bruggeman J, Ané C (2017) Rphylopars: Fast multivariate phylogenetic com- parative methods for missing data and within-species variation. Methods in Ecology and Evolution 8(1): 22-27. https://doi.org/10.1111/2041-210X.12612 Greilhuber J, Dolezel J, Lysik MA, Bennett MD (2005) The origin, evolution and proposed stabilization of the terms “genome size” and “C-value” to describe nuclear DNA contents. Annals of Botany 95(1): 255-260. https://doi.org/10.1093/aob/mci019 Guo W-Y, van Kleunen M, Pierce S, Dawson W, Essl F, Kreft H, Maurel N, Pergl J, Seebens H, Weigelt P, Pysek P (2019) Domestic gardens play a dominant role in selecting alien species with adaptive strategies that facilitate naturalization. Global Ecology and Biogeography 28(5): 628-639. https://doi.org/10.1111/geb.12882 Haider S, Alexander J, Dietz H, Trep] L, Edwards PJ, Kueffer C (2010) The role of bioclimatic origin, residence time and habitat context in shaping non-native plant distributions along an altitudinal gradient. Biological Invasions 12(12): 4003-4018. https://doi.org/10.1007/ s10530-010-9815-7 Inouye DW (2020) Effects of climate change on alpine plants and their pollinators. Annals of the New York Academy of Sciences 1469(1): 26-37. https://doi.org/10.1111/nyas.14104 Iseli E, Chisholm C, Lenoir J, Haider S, Seipel T, Barros A, Hargreaves AL, Kardol P, Lembre- chts JJ, McDougall K, Rashid I, Rumpf SB, Arévalo JR, Cavieres L, Daehler C, Dar PA, Endress B, Jakobs G, Jiménez A, Kiiffer C, Mihoc M, Milbau A, Morgan JW, Naylor BJ, Pauchard A, Ratier Backes A, Reshi ZA, Rew LJ, Righetti D, Shannon JM, Valencia G, Walsh N, Wright GT, Alexander JM (2023) Rapid upwards spread of non-native plants in mountains across continents. Nature Ecology & Evolution 7(3): 405-413. https://doi. org/10.1038/s41559-022-01979-6 Jin Y, Qian H (2019) V.PhyloMaker: An R package that can generate very large phylogenies for vascular plants. Ecography 42(8): 1353-1359. https://doi.org/10.1111/ecog.04434 Kindt R (2020) WorldFlora: An R package for exact and fuzzy matching of plant names against the World Flora Online taxonomic backbone data. Applications in Plant Sciences 8(9): e11388. https://doi.org/10.1002/aps3.11388 Kleyer M, Bekker RM, Knevel IC, Bakker JP, Thompson K, Sonnenschein M, Poschlod P, Van Groenendael JM, Klimes L, Klimesova J, Klotz S$, Rusch GM, Hermy M, Adriaens D, Boedeltje G, Bossuyt B, Dannemann A, Endels P, Gétzenberger L, Hodgson JG, Jackel AK, Kiihn I, Kunzmann D, Ozinga WA, Rémermann C, Stadler M, Schlegelmilch J, Steendam HJ, Tackenberg O, Wilmann B, Cornelissen JHC, Eriksson O, Garnier E, Peco B (2008) The LEDA Traitbase: A database of life-history traits of the Northwest European flora. Journal of Ecology 96(6): 1266-1274. https://doi.org/10.1111/j.1365- 2745.2008.01430.x Knight CA, Molinari NA, Petrov DA (2005) The large genome constraint hypothesis: Evolu- tion, ecology and phenotype. Annals of Botany 95(1): 177-190. https://doi.org/10.1093/ aob/mci011 Koide D, Yoshida K, Daehler CC, Mueller-Dombois D (2017) An upward elevation shift of native and non-native vascular plants over 40 years on the island of Hawai'i. Journal of Vegetation Science 28(5): 939-950. https://doi.org/10.1111/jvs.12549 Clonal species invade larger and faster 43 KubeSova M, Moravcova L, Suda J, Jarosik V, Pysek P (2010) Naturalized plants have smaller genomes than their non-invading relatives: A flow cytometric analysis of the Czech alien flora. Preslia 82: 81—96. Liu J, Dong M, Miao SL, Li ZY, Song MH, Wang RQ (2006) Invasive alien plants in China: Role of clonality and geographical origin. Biological Invasions 8(7): 1461-1470. https://doi.org/10.1007/s10530-005-5838-x Liu YY, Sun Y, Miiller-Scharer H, Yan R, Zhou ZX, Wang YJ, Yu FH (2019) Do invasive alien plants differ from non-invasives in dominance and nitrogen uptake in response to varia- tion of abiotic and biotic environments under global anthropogenic change? The Science of the Total Environment 672: 634-642. https://doi.org/10.1016/j.scitotenv.2019.04.024 Lopes S, Mota L, Castro M, Nobre G, Novoa A, Richardson DM, Loureiro J, Castro S (2021) Genome size variation in Cactaceae and its relationship with invasiveness and seed traits. Biological Invasions 23(10): 3047-3062. https://doi.org/10.1007/s10530-021-02557-w Marini L, Bertolli A, Bona E, Federici G, Martini EF, Prosser F Bommarco R (2013) Beta-diver- sity patterns elucidate mechanisms of alien plant invasion in mountains. Global Ecology and Biogeography 22(4): 450-460. https://doi.org/10.1111/geb.12006 McDougall KL, Khuroo AA, Loope LL, Parks CG, Pauchard A, Reshi ZA, Rushworth I, Kueffer C (2011) Plant invasions in mountains: Global lessons for better management. Mountain Research and Development 31(4): 380-387. https://doi.org/10.1659/MRD- JOURNAL-D-11-00082.1 Menzel A, Hempel S, Klotz S, Moora M, PySek P, Rillig MC, Zobel M, Kithn I (2017) Mycor- rhizal status helps explain invasion success of alien plant species. Ecology 98(1): 92-102. https://doi.org/10.1002/ecy.1621 Moodley D, Geerts S, Richardson DM, Wilson JRU (2013) Different traits determine intro- duction, naturalization and invasion success in woody plants: Proteaceae as a test case. PLOS ONE 8(9): e75078. https://doi.org/10.1371/journal.pone.0075078 Pauchard A, Kueffer C, Dietz H, Daehler CC, Alexander J, Edwards PJ, Arévalo JR, Cavieres LA, Guisan A, Haider S, Jakobs G, McDougall K, Millar CI, Naylor BJ, Parks CG, Rew LJ, Seipel T (2009) Aint no mountain high enough: Plant invasions reaching new elevations. Frontiers in Ecology and the Environment 7(9): 479-486. https://doi. org/10.1890/080072 Pellicer J, Leitch IJ (2020) The Plant DNA C-values database (release 7.1): An updated online repository of plant genome size data for comparative studies. The New Phytologist 226(2): 301-305. https://doi.org/10.1111/nph.16261 Pergl J, Pysek P, Bacher S, Essl EK Genovesi P, Harrower CA, Hulme PE, Jeschke JM, Kenis M, Kiihn I, Perglova I, Rabitsch W, Roques A, Roy DB, Roy HE, Vila M, Winter M, Nentwig W (2017) Troubling travellers: Are ecologically harmful alien species associated with particular introduction pathways? NeoBiota 32: 1-20. https://doi.org/10.3897/neo- biota.32.10199 Prinzing A, Durka W, Klotz S, Brandl R (2001) The niche of higher plants: Evidence for phy- logenetic conservatism. Proceedings of the Royal Society B: Biological Sciences 268(1483): 2383-2389. https://doi.org/10.1098/rspb.2001.1801 44 Miao-Miao Zheng et al. / NeoBiota 91: 29-48 (2024) PySek P (1997) Clonality and plant invasions: can a trait make a difference? In: de Kroon H, van Groenendael J (Eds) Ecology and evolution of clonal plants. Backhuys Publishers, Leiden, 405-427. Pysek P, Jarosik V, Pergl J, Randall R, Chytry M, Kithn I, Tichy L, Danihelka J, Chrtek Jun J, Sadlo J (2009a) The global invasion success of Central European plants is related to dis- tribution characteristics in their native range and species traits. Diversity & Distributions 15(5): 891-903. hteps://doi.org/10.1111/j.1472-4642.2009.00602.x Pysek P, Krivanek M, Jarosik V (2009b) Planting intensity, residence time, and species traits determine invasion success of alien woody species. Ecology 90: 2734-2744. https://doi. org/10.1890/08-0857.1 Pysek P, Jarosik V, Pergl J, Wild J (2011) Colonization of high altitudes by alien plants over the last two centuries. Proceedings of the National Academy of Sciences of the United States of America 108(2): 439-440. https://doi.org/10.1073/pnas.1017682108 PySek P, Danihelka J, Sadlo J, Chrtek Jr JC, Chytry M, Jarosik V, Kaplan Z, Krahulec EK Morav- cova L, Pergl J, Stajerova K, Tichy L (2012) Catalogue of alien plants of the Czech Republic (2"4 edn.): Checklist update, taxonomic diversity and invasion patterns. Preslia 84: 155-255. Pysek P, Manceur AM, Alba C, McGregor KE, Pergl J, Stajerova K, Chytry M, Danihelka J, Kartesz J, KlimeSova J, Lucanova M, Moravcova L, Nishino M, Sadlo J, Suda J, Tichy L, Kihn I (2015) Naturalization of central European plants in North America: Species traits, habitats, propagule pressure, residence time. Ecology 96(3): 762-774. https://doi. org/10.1890/14-1005.1 Pysek P, Pergl J, Ess] E Lenzner B, Dawson W, Kreft H, Weigelt P, Winter M, Kartesz J, Nishino M, Antonova LA, Barcelona JE Cabezas FJ, Cardenas D, Cardenas-Toro J, Castafio N, Chacon E, Chatelain C, Dullinger S, Ebel AL, Figueiredo E, Fuentes N, Genovesi P, Groom QJ, Henderson L, Inderjit N, Kupriyanov A, Masciadri S, Maurel N, Meerman J, Morozova O, Moser D, Nickrent DL, Nowak PM, Pagad S, Patzelt A, Pelser PB, Seebens H, Shu WS, Thomas J, Velayos M, Weber E, Wieringa JJ, Baptiste MP, van Kleunen M (2017) Naturalized alien flora of the world: Species diversity, taxonomic and phylogenetic patterns, geographic distribution and global hotspots of plant invasion. Preslia 89(3): 203- 274. https://doi.org/10.23855/preslia.2017.203 Pysek P, Guo W-Y, Stajerova K, Moora M, Bueno CG, Dawson W, Essl F, Gerz M, Kreft H, Pergl J, van Kleunen M, Weigelt P, Winter M, Zobel M (2019) Facultative mycor- rhizal associations promote plant naturalization worldwide. Ecosphere 10(11): e02937. https://doi.org/10.1002/ecs2.2937 Pyek P, Sadlo J, Chrtek J, Chytry M, Kaplan Z, Pergl J, Pokorna A, Axmanova I, Cuda J, Dolezal J, Dievojan P, Hejda M, Koéar P, Kortz A, Lososova Z, Lustyk P, Skalova H, Stajerova K, Veteta M, Danihelka J (2022) Catalogue of alien plants of the Czech Republic (3 edn.): species richness, status, distributions, habitats, regional invasion levels, introduction path- ways and impacts. Preslia 94: 447-577. https://doi.org/10.23855/preslia.2022.447 Pysek P, Lu¢éanova M, Dawson W, Essl F, Kreft H, Leitch Jj, Lenzner B, Meyerson LA, Pergl J, van Kleunen M, Weigelt P, Winter M, Guo W-Y (2023) Small genome size and variation in ploidy levels support the naturalization of vascular plants but constrain their invasive spread. The New Phytologist 239(6): 2389-2403. https://doi.org/10.1111/nph.19135 Clonal species invade larger and faster 45 R Core Team (2023) R: The R Project for Statistical Computing. Rahbek C, Borregaard MK, Antonelli A, Colwell RK, Holt BG, Nogues-Bravo D, Rasmussen CM®@, Richardson K, Rosing MT, Whittaker RJ, Fjeldsa J (2019) Building mountain biodiversity: Geological and evolutionary processes. Science 365(6458): 1114-1119. https://doi.org/10.1126/science.aax0151 Rangel TE, Edwards NR, Holden PB, Diniz-Filho JAF, Gosling WD, Coelho MTP, Cassemiro FAS, Rahbek C, Colwell RK (2018) Modeling the ecology and evolution of biodiversity: Biogeographical cradles, museums, and graves. Science 361(6399): eaar5452. https://doi. org/10.1126/science.aar5452 Revell LJ (2012) phytools: An R package for phylogenetic comparative biology (and other things). Methods in Ecology and Evolution 3(2): 217-223. https://doi.org/10.1111/ j-2041-210X.2011.00169.x Schielzeth H (2010) Simple means to improve the interpretability of regression coefficients. Methods in Ecology and Evolution 1(2): 103-113. https://doi.org/10.1111/j.2041- 210X.2010.00012.x Si L, Ho T, Ane C, Lachlan R, Tarpinian K, Feldman R, Yu Q, Van Der Bijl W, Maspons J, Vos R, Lam M, Ho ST (2022) Package “phylolm.” Skalova H, Guo W-Y, Wild J, PySek P (2017) Ambrosia artemisiifolia in the Czech Republic: History of invasion, current distribution and prediction of future spread. Preslia 89(1): 1-16. https://doi.org/10.23855/preslia.2017.001 Soudzilovskaia NA, He J, Rahimlou S, Abarenkov K, Brundrett MC, Tedersoo L (2022) Fun- galRoot v.2.0 — an empirical database of plant mycorrhizal traits. The New Phytologist 235(5): 1689-1691. https://doi.org/10.1111/nph.18207 Suda J, Meyerson LA, Leitch IJ, Pysek P (2015) The hidden side of plant invasions: The role of genome size. [he New Phytologist 205(3): 994-1007. https://doi.org/10.1111/nph.13107 Tichy L, Axmanova I, Dengler J, Guarino R, Jansen F, Midolo G, Nobis MP, Meerbeek KV, Adié S, Attorre F, Bergmeier E, Biurrun I, Bonari G, Bruelheide H, Campos JA, Carni A, Chiarucci A, Cuk M, CuSterevska R, Didukh Y, Dité D, Dité Z, Dziuba T, Fanelli G, Fernandez-Pascual E, Garbolino E, Gavilan RG, Gégout J-C, Graf U, Giiler B, Hajek M, Hennekens SM, Jandt U, Jaskova A, Jiménez-Alfaro B, Julve P, Kambach S, Karger DN, Karrer G, Kavgaci A, Knollova I, Kuzemko A, Kiizmi¢ F, Landucci FE Lengyel A, Lenoir J, Marcend C, Moeslund JE, Novak P, Pérez-Haase A, Peterka T, Pielech R, Pignatti A, Rasomavicius V, Risina S, Saatkamp A, Silc U, Skvorc Z, Theurillat J-P Wohlgemuth T, Chytry M (2022) Ellenberg-type indicator values for European vascular plant species. Journal of Vegetation Science 34(1): 13168. https://doi.org/10.1111/ jvs.13168 Tung Ho LS, Ané C (2014) A linear-time algorithm for Gaussian and non-Gaussian trait evolu- tion models. Systematic Biology 63(3): 397-408. https://doi.org/10.1093/sysbio/syu005 van Kleunen M, Weber E, Fischer M (2010) A meta-analysis of trait differences between invasive and non-invasive plant species. Ecology Letters 13(2): 235-245. https://doi. org/10.1111/j.1461-0248.2009.01418.x van Kleunen M, Dawson W, Essl F, Pergl J, Winter M, Weber E, Kreft H, Weigelt P, Kartesz J, Nishino M, Antonova LA, Barcelona JE Cabezas FJ, Cardenas D, Cardenas-Toro J, 46 Miao-Miao Zheng et al. / NeoBiota 91: 29-48 (2024) Castaho N, Chacon E, Chatelain C, Ebel AL, Figueiredo E, Fuentes N, Groom QJ, Henderson L, Inderjit, Kupriyanov A, Masciadri S$, Meerman J, Morozova O, Moser D, Nickrent DL, Patzelt A, Pelser PB, Baptiste MP, Poopath M, Schulze M, Seebens H, Shu W, Thomas J, Velayos M, Wieringa JJ, PySek P (2015) Global exchange and accumulation of non-native plants. Nature 525(7567): 100-103. https://doi.org/10.1038/nature14910 van Kleunen M, Essl F, Pergl J, Brundu G, Carboni M, Dullinger S, Early R, Gonzalez-Moreno P, Groom QJ, Hulme PE, Kueffer C, Kitihn I, Maguas C, Maurel N, Novoa A, Parepa M, PySek PB, Seebens H, Tanner R, Touza J, Verbrugge L, Weber E, Dawson W, Kreft H, Weigelt P, Winter M, Klonner G, Talluto MV, Dehnen-Schmutz K (2018) The changing role of ornamental horticulture in alien plant invasions. Biological Reviews of the Cam- bridge Philosophical Society 93(3): 1421-1437. https://doi.org/10.1111/brv.12402 van Kleunen M, Pysek P, Dawson W, Essl F, Kreft H, Pergl J, Weigelt P, Stein A, Dullinger S, K6nig C, Lenzner B, Maurel N, Moser D, Seebens H, Kartesz J, Nishino M, Aleksanyan A, Ansong M, Antonova LA, Barcelona JE, Breckle SW, Brundu G, Cabezas FJ, Cardenas D, Cardenas-Toro J, Castafio N, Chacén E, Chatelain C, Conn B, de SA Dechoum M, Dufour-Dror JM, Ebel AL, Figueiredo E, Fragman-Sapir O, Fuentes N, Groom QJ, Henderson L, Inderjit, Jogan N, Krestov P, Kupriyanov A, Masciadri S, Meerman J, Morozova O, Nickrent D, Nowak A, Patzelt A, Pelser PB, Shu W, Thomas J, Uludag A, Velayos M, Verkhosina A, Villasehor JL, Weber E, Wieringa JJ, Yazlik A, Zeddam A, Zykova E, Winter M (2019) The Global Naturalized Alien Flora (GloNAF) database. Ecology 100(1): e02542. https://doi.org/10.1002/ecy.2542 van Kleunen M, Xu X, Yang Q, Maurel N, Zhang Z, Dawson W, Essl FE, Kreft H, Pergl J, PySek P, Weigelt P, Moser D, Lenzner B, Fristoe TS (2020) Economic use of plants is key to their naturalization success. Nature Communications 11(1): 1-12. https://doi.org/10.1038/ s41467-020-16982-3 Wang YJ, Chen D, Yan R, Yu FH, van Kleunen M (2019) Invasive alien clonal plants are competitively superior over co-occurring native clonal plants. Perspectives in Plant Ecol- ogy, Evolution and Systematics 40: 125484. https://doi.org/10.1016/j.ppees.2019.125484 Wang YJ, Liu YY, Chen D, Du DL, Miiller-Scharer H, Yu FH (2024) Clonal functional traits favor the invasive success of alien plants into native communities. Ecological Applications 34(1): e2756. https://doi.org/10.1002/eap.2756 Wang H, Liu H, Cao G, Ma Z, Li Y, Zhang FE, Zhao X, Zhao X, Jiang L, Sanders NJ, Classen AT, He J-S (2020) Alpine grassland plants grow earlier and faster but biomass remains unchanged over 35 years of climate change. Ecology Letters 23: 701-710. https://doi. org/10.1111/ele.13474 Williamson M, PySek P, Jarosik V, Prach K (2005) On the rates and patterns of spread of alien plants in the Czech Republic, Britain, and Ireland. Ecoscience 12(3): 424-433. https://doi. org/10.2980/i1195-6860-12-3-424.1 Wolf A, Zimmerman NB, Anderegg WRL, Busby PE, Christensen J (2016) Altitudinal shifts of the native and introduced flora of California in the context of 20"-century warming. Global Ecology and Biogeography 25(4): 418-429. https://doi.org/10.1111/ geb. 12423 Clonal species invade larger and faster 47 Appendix | e CY acne e e at seg 00° eon ee gs ens ENS per ant 208 Sg coors? gy vd geet 900 4 \] wn 0.53 | -0.42 |-0.51|-0.63 | 0.23 10.46 | aex 3005 pen fm ; . : se % a Ane jem jo | oe is i i a rave) Correlation 1.0 vy 0.37 oF z = re) 9 Ee me we Figure Al. Pearson’s correlation for all continuous variables collected in the study. EIE_range, elevation range; INV_speed, invasion speed; FLOW_time, flowering time; EIV_L, Ellenberg indicator values of light; EIV_T, Ellenberg indicator values of temperature; EIV_N, Ellenberg indicator values of nutri- ents; NAT_range, native range; NATLZ_range, naturalized range; Ecouse, economic use sum; Cx_value, monoploid genome size; MRT, minimum residence time. 48 Miao-Miao Zheng et al. / NeoBiota 91: 29-48 (2024) (a) (b) (c) 900 = f 900 - 900 - é | () Pimean = 746.29 2 Ta © = = 600 = 600- 600 - 2 | = © > 2 lw 300 = 300- 300 - Accidental Deliberate (n = 31) (n= 27) (e) r 8- 8- ay © © 2 = «- 6- xe} ® ® 2 7) < ors 4- 7) © > £ 2- oh Deliberate (n= 22) Life Form MYC Pathway Figure A2. Violin plots of the elevation ranges and invasion speeds for the 58 alien species consider- ing their life spans, mycorrhizal types, and introduction pathways. For the boxplots inside each violin plot, the horizontal line represents the median, the red dot indicates the mean, and the box represents the interquartile range. P-values were calculated using phylogenetic ANOVA models, and none of them are significant. Supplementary material | Data used for the analysis Authors: Miao-Miao Zheng, Petr Pysek, Kun Guo, Hasigerili, Wen-Yong Guo Data type: csv Copyright notice: This dataset is made available under the Open Database License (http://opendatacommons.org/licenses/odbl/1.0/). The Open Database License (ODbL) is a license agreement intended to allow users to freely share, modify, and use this Dataset while maintaining this same freedom for others, provided that the original source and author(s) are credited. Link: https://doi.org/10.3897/neobiota.91.115675.suppl1