Research Article |
Corresponding author: Wen-Yong Guo ( guowyhgy@gmail.com ) Academic editor: Gerhard Karrer
© 2024 Miao-Miao Zheng, Petr Pyšek, Kun Guo, Hasigerili, Wen-Yong Guo.
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.
Citation:
Zheng M-M, Pyšek P, 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
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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. However, how quickly alien species spread upward and what drives their invasion remains insufficiently understood. 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 underlying factors driving these variables. We collected species traits relevant for invasion (e.g., clonality, flowering 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 spatiotemporal dynamics of alien species in high-altitude ecosystems and thus employing apposite measures to reduce the threat to native plant species.
Alien plant species, clonal trait, elevation range, human activities, invasion speed, mountain invasions
Mountains are of crucial importance for biodiversity conservation (
In mountainous areas, the influx of alien species typically commences at lower elevations before gradually spreading upward over time (
In addition to the minimum residence time, various characteristics of alien species, e.g., introduction pathways (
Karyological characteristics were recently suggested as an important trait underpinning plant invasion success, and species with small genomes proved to be at an advantage in the process of alien plants’ naturalization (
While the magnitude of a species’ native range significantly influences its preadaptation to the introduced environment, and a broader native range generally fosters greater readiness for the establishment and thriving within the novel environment (
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 characteristics of these plants, including both species’ inherent traits (e.g., clonality, flowering 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 drivers 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 species, and play distinct roles in determining the elevational range (and changes) and invasion speed.
Species data were obtained from an existing dataset (
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 maximum elevation year, due to the possibility of inadequate sampling; and negative values. We used the ‘WorldFlora’ R package (
We obtained data on the clonality (modular species with potential vegetative reproduction or unitary species without this potential), flowering time (as the first month of flowering), life span (annual, perennial, or both), and invasion status (casual, naturalized, invasive) of the 58 alien species from existing datasets (
The height of each species was extracted from the LEDA database (https://uol.de/en/landeco/research/leda) (
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 data were obtained from the GloNAF (Global Naturalized Alien Flora) database (
Information on the introduction pathway (deliberate or accidental introduction) of each alien species to the Czech Republic was taken from
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 (
Given the presence of missing data for certain species, we undertook gap-filling processes as follows. For the 20 species lacking monoploid genome size, imputation was carried out utilizing the full dataset of 12,273 species (
All data analyses and visualizations were performed in R v4.2.1 (
The elevation ranges of the 58 alien species varied from 132 m to 1095 m (Fig.
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.
Both elevation range and invasion speed differed with regard to clonality (Fig.
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).
Phylogenetic regressions revealed different factors driving the elevation range and invasion speed for clonal and non-clonal species (Figs
Standardized estimates and associated 95% confidence intervals obtained from phylogenetic regressions 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).
Relationships between elevation ranges and important variables identified from phylogenetic regressions (See Fig.
Using more than 26,000 historical occurrence records of 58 alien plant species introduced to the Czech Republic after 1738, we found a substantial upward shift along the elevations for the majority of the species analyzed (Fig.
Among the various factors tested in this study, alien species with clonal reproduction exhibited not only significantly broader elevation ranges but also faster upward expansion compared to their non-clonal counterparts (Fig.
For other traits tested with regard to the elevation expansion of alien species, we observed a negative association between monoploid genome size and elevation range for clonal aliens (Fig.
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.
It is gradually recognized that the position on the introduction-naturalization-invasion continuum is a good indicator of species’ minimum residence time (
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 (
Although we considered several factors related to species’ expansion along the elevation gradient, several important variables, such as climate and soil properties, were missing from the analysis. It appears necessary to include these variables in future studies 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 particular, 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’ 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.
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).
Pearson’s correlation for all continuous variables collected in the study. ElE_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 nutrients; NAT_range, native range; NATLZ_range, naturalized range; Ecouse, economic use sum; Cx_value, monoploid genome size; MRT, minimum residence time.
Violin plots of the elevation ranges and invasion speeds for the 58 alien species considering 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.
Data used for the analysis
Data type: csv