Research Article |
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Corresponding author: Kristine Bakke Westergaard ( kristine.b.westergaard@ntnu.no ) Academic editor: Zarah Pattison
© 2025 Tor Henrik Ulsted, Kristine Bakke Westergaard, Wayne Dawson, James D. M. Speed.
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:
Ulsted TH, Westergaard KB, Dawson W, Speed JDM (2025) Horizon scanning of potential new alien vascular plant species and their climatic niche space across the Arctic. NeoBiota 104: 1-26. https://doi.org/10.3897/neobiota.104.165054
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The terrestrial Arctic faces increasing vulnerability to alien plant invasions due to climate change and intensifying human activities. Using a data-driven horizon scanning approach that leverages the Global Naturalized Alien Flora (GloNAF) database, species occurrence data from the Global Biodiversity Information Facility (GBIF), and climate data from WorldClim, we identified 2,554 potential new alien vascular plant species with climatic niches overlapping Arctic floristic provinces. Six major potential hotspots for introductions were detected, with western Alaska, southwestern and southeastern Greenland, northern Iceland, Fennoscandia, and Kanin–Pechora showing the highest numbers of potential alien species. Potential source regions for these species extended globally across diverse climate zones, with substantial contributions from proximate temperate regions in Europe and North America. Taxonomic analysis revealed that most Arctic floristic provinces exhibited compositions similar to global patterns, with only Franz Joseph Land showing significant deviation after multiple comparison corrections, although island provinces generally demonstrated greater compositional distinctiveness than mainland provinces. Zero-inflated beta regression analysis confirmed our hypothesis that species with higher absolute latitude distributions demonstrate greater potential for climatic overlap with Arctic floristic provinces. Our findings emphasize the need to develop effective biosecurity measures in high-risk regions and to proactively manage emerging invasion risks across the rapidly changing terrestrial Arctic ecosystems. This will provide a foundation for supporting community-based monitoring networks essential for early detection and rapid response initiatives.
Arctic biodiversity, bioclimatic modelling, climate change, early detection rapid response, horizon scanning, invasion science
The Arctic region faces escalating threats from biological invasions driven by rising human activity and climate change (
A comprehensive inventory of alien plant taxa in the Arctic previously documented 341 taxa, providing a valuable foundation for understanding patterns of establishment and spread across the region. Of these, 188 had become naturalized in at least one floristic province (i.e., subdivisions of the Arctic based on floristic differences;
Early detection and rapid response (EDRR) is recognized as a cost-effective strategy for managing alien taxa (e.g.,
Horizon scanning is a systematic process for identifying future threats and opportunities to inform proactive management (
Here we employ a data-driven horizon scanning approach to identify potential new alien vascular plants across the terrestrial Arctic. Although Arctic plant habitats form a diverse and complex mosaic, regional east–west patterns of vascular plant species richness reflect differences in glaciation history, geology, and geography (i.e., floristic provinces;
We used the delimitation of the terrestrial Arctic provided by the Circumpolar Arctic Vegetation Map (CAVM;
We used the Global Naturalized Alien Flora database (GloNAF;
Workflow diagram illustrating the data processing and filtering steps to identify plant taxa from the Global Naturalized Alien Flora (GloNAF) database that are absent from the Arctic region. The process consists of four main stages: (1) data wrangling of inventory lists, including the Arctic Plant Inventory, Arctic Alien Plant List, and GloNAF database; (2) standardization through synonym checking; (3) combining and filtering of standardized lists to remove duplicates and identify present and absent taxa; and (4) occurrence data refinement through the removal of infraspecific taxa. The workflow begins with 13,939 alien naturalized taxa from GloNAF and concludes with 11,482 filtered species with occurrence records. Color coding indicates downloaded data (green), supplementary data (light orange), taxa addition steps (blue), removal steps (pink), and indicators for descriptions (light blue). Numbers in parentheses represent the count of taxa at each processing stage.
The GloNAF list contained 13,939 vascular plant taxa known to be naturalized or alien worldwide. Both “naturalized” (has established self-sustaining populations in the wild) and “alien” (when it is not clear whether they are naturalized) taxa were included in the filtering process. The Arctic Plant Inventory List comprises 2,065 taxa, while the Arctic Alien Plant List contains 334 taxa. Both Arctic lists include taxon-specific distribution information on presence and absence in the 23 floristic provinces of the Arctic. First, the native taxa on the Arctic Plant Inventory List were sorted into “Inventory Arctic present” (including “frequent,” “rare,” “scattered,” and “present” taxa) and “Inventory Arctic absent” (including “casual,” “absent,” “borderline,” and “unknown” taxa; Fig.
We standardized scientific names for each list using the World Flora Online (WFO) package in R (
We used the name_backbone function in the R package rgbif (
These usage keys were then used with the occ_download function (Derived dataset GBIF.org 2025b) to retrieve records with status “present,” excluding records with geospatial issues (an ensemble of 30 different issues related to coordinates, elevation, and depth;
We filtered out taxa where the logarithm of the number of occurrence records was fewer than the number of climate dimensions used to calculate hypervolume space (four; see “climate space” below). A hypervolume is a set of points within an n-dimensional space that represents the ecological niche of a taxon, where each axis corresponds to a biologically relevant variable. This framework ensures flexible modeling of complex, high-dimensional data distributions (
Climate data were downloaded from WorldClim 2.0 (
The climate data were then cropped to the Arctic as delineated by CAVM, and the Pearson correlation coefficient (|rp|) was employed as a statistical measure to evaluate pairwise linear covariations between the 19 bioclimatic variables. The absolute value of the Pearson correlation coefficient was used to quantify the strength of the correlation, irrespective of its direction. This approach allowed us to identify and select bioclimatic variables that effectively characterized the Arctic climate while ensuring these variables did not exhibit high multicollinearity (|rp| < 0.5; Suppl. material
Each taxon’s longitude and latitude coordinates were cleaned using the coordinateCleaner (
We used an adapted version of a hypervolume approach (
The climatic niche space was constructed using kernel density estimation (KDE) with the hypervolume package in R (
where m is the number of occurrences, n is the dimensionality, and X is the data vector in each dimension. This corresponds to the Silverman rule of thumb for multivariate data (
Next, each taxon underwent two exclusion tests to limit the time consumed by the analysis process (Suppl. material
The same inclusion analysis method was then applied spatially to create projection maps. We used the accurate algorithm, which estimates the probability density at each test point and excludes points below a threshold of 0.5. Although this method is slower, it provides more reliable results than the fast algorithm (
The probability analysis calculates the probability density at various points within or outside the taxon hypervolume. This is done by computing a weighted sum of the probability densities of all subsampled random points in the hypervolume, where the weights are proportional to the distance from the test point raised to the power of −1 (inverse distance weighting;
To validate the model used in the climate niche analysis, we applied the same process to the vascular plant taxa already present in the Arctic as listed in the Arctic Plant Inventory List and Arctic Alien Plant List. This included 1,558 known Arctic taxa downloaded from GBIF using the occ_download function in the R package rgbif (
We conducted several analyses to characterize potential new alien vascular plant taxa distributions and patterns. First, we quantified each taxon’s potential total area of occupancy by applying the hypervolume project function from the hypervolume package in R (
where n is the unique number of potential new alien taxa in each order in each floristic province, and N is the total number of potential new alien taxa in each floristic province. This analysis revealed how taxonomic composition varies across the Arctic. Furthermore, we analyzed the statistical significance of taxonomic compositional differences between Arctic floristic provinces and the global baseline (GloNAF) using chi-square goodness-of-fit tests. The standard asymptotic assumptions of the chi-square test are often violated in ecological data due to low expected frequencies in many categories. To address this, we employed a Monte Carlo simulation with 2,000 replicates (B = 2,000) to calculate robust p-values empirically (
where χ2 is the chi-square statistic, n is the sample size (total number of taxa) in each floristic province, and k is the number of taxonomic orders. Effect sizes were interpreted as small (V < 0.3), medium (0.3 ≤ V ≤ 0.5), or large (V > 0.5) differences from the global baseline composition (GloNAF) (
To investigate the potential source regions of future alien plants in the Arctic, we first standardized the geographical representation of taxon occurrences across regions by calculating the centroid (mean latitude and longitude) of all occurrence points for each taxon within each botanical country (Level 3;
Finally, to test whether taxa originating from higher latitudes tend to have greater climatic overlap with the Arctic, we modeled climate overlap as a function of the taxa’s absolute median latitude using a zero-inflated beta regression model in R (BEZI family in GAMLSS;
We compared models using the Akaike Information Criterion (AIC) and Bayesian Information Criterion (BIC) to determine which best explained the relationship between latitude and climate overlap patterns. AIC was used to identify models with strong predictive accuracy, while BIC—with its stronger penalty for complexity—helped ensure model parsimony. Using both criteria provided complementary perspectives on model selection (Suppl. material
The climatic niche overlap analysis identified 2,554 vascular plant taxa as potential new alien taxa to the Arctic (Suppl. material
The potential richness of new alien plants (i.e., the number of species with suitable climate at a given location across the Arctic) ranged from 0 to 1,273 species per km2 cell (Fig.
Hotspots of potential new alien vascular plant species richness across the Arctic, based on 1 × 1 km resolution models of GloNAF species with climatic niches overlapping Arctic regions. Darker colors represent lower numbers of potential alien species, while brighter colors represent higher numbers. The terrestrial Arctic is defined by the extent of the colored areas, delimited according to the Circumpolar Arctic Vegetation Map (CAVM;
The top five potential new alien species having the largest projected areas of climatic suitability in the Arctic are presented in Fig.
Potential distribution of the top five potential new alien vascular plant species in the Arctic. Left panel: potential area of occupancy (regardless of probability). Red areas indicate potential presence, while black areas indicate no potential occupancy. Right panel: potential climatic suitability. Warmer colors (yellow to red) indicate higher climatic suitability for the species.
To assess whether the taxonomic composition of the potential new alien flora differed from global patterns, we compared the relative richness of plant orders in each floristic province to the GloNAF baseline (Fig.
Taxonomic composition of potential new alien taxa across Arctic floristic provinces. The bars represent the relative richness of each taxonomic order within each floristic province, sorted from west (left) to east (right), and include the comparison with the GloNAF database as a separate bar on the right side. Different colors represent different taxonomic orders.
The potential new alien species identified in this study originate from a wide range of botanical countries worldwide (Fig.
Potential source regions for new alien taxa to the Arctic. Green points indicate species centroids within botanical countries around the world (based on the World Geographical Scheme for Recording Plant Distributions), representing potential source regions. The blue area outlines the Arctic.
Both AIC and BIC consistently identified the full model as the best-fitting, providing strong support for the inclusion of latitude as a predictor of all three components: the mean climate overlap, its variance, and the probability of zero overlap (Suppl. material
The relationship between taxon's absolute median latitude and Arctic niche overlap presented in two panels. Panel A (top) shows the expected overlap magnitude given latitude. Gray points represent individual species data, while the green line represents the conditional expected overlap when overlap exceeds zero (E (Overlap | Overlap > 0)), and the orange line shows the combined expectation. Panel B (bottom) displays the probability of any Arctic niche overlap as a function of latitude, shown by the blue curve. Together, these panels demonstrate that both the probability of Arctic niche overlap and its expected magnitude increase with species’ absolute median latitude, with the probability approaching 100% at the highest latitudes (> 60°).
Our study identifies 2,554 potential new alien vascular plants that have a climatic niche overlap in the Arctic. We reveal clear spatial patterns in climate suitability across Arctic floristic provinces and locate potential hotspots in southern regions, including western Alaska and northern Fennoscandia. The majority of species displayed limited predicted occupancy, with only 42 species exceeding 20% potential coverage of the terrestrial Arctic region. The highest predicted occupancies were found among four species that—following the precautionary principle and taxonomic filtering criteria—already have recorded occurrence points for one or several infraspecific taxa within the Arctic. Additionally, species currently established in adjacent biogeographic zones, such as Senecio nemorensis L. (62% occupancy) and Thalictrum foetidum L. (51%), showed high potential climate suitability in the Arctic (Derived dataset GBIF.org 2025b). This finding is consistent with earlier studies showing that proximity to the Arctic enhances the likelihood of successful establishment (
Compiling a list of relevant taxa for horizon scanning involves gathering species occurrence data and their alien status and harmonizing inconsistencies in taxonomy and nomenclature from different data providers and sources varying in accuracy and age. Our data-driven workflow effectively collates and mass-curates digital taxon data from site-specific taxon lists and databases, employs a taxonomic backbone to harmonize nomenclature across data providers, and flags cases for manual review. Through the climate suitability analysis, we then generate a prioritized list of relevant potential new alien taxa to the Arctic based on macroclimatic niche overlap, providing a foundation for further assessment of their introduction pathways, invasion risk, and ecological impacts.
Of the top five potential new alien species in the Arctic (Fig.
As
Several species with known Arctic distributions belong to taxonomic complexes with multiple subspecies, some or all of which are known to be naturalized elsewhere in the world according to GloNAF (
The potential source regions for new alien plant species in the Arctic extend globally across nearly all climate zones (Fig.
Such widespread source geography and composition present significant challenges to biosecurity efforts, as they imply that preventive measures cannot focus exclusively on particular geographic pathways. As identified by
The diversity of source regions suggests a complex interplay between plant traits and human-mediated factors in determining Arctic introduction potential. While physiological traits related to cold stress adaptations, competitive ability, dispersal limitations, and microclimate requirements may currently limit establishment (
While our climatic niche horizon scanning approach allowed analysis of extensive datasets compared to expert consensus methods, we acknowledge several considerations that inform future research directions. Taxonomic standardization challenges included the necessary collapsing of certain infraspecific taxa into species-level classifications, which contributed to some apparently novel potential new alien candidates already having conspecific populations in the Arctic. Occasional misclassifications or errors in data labeling are possible (e.g., Hylotelephium maximum (L.) Holub had an erroneous record in the Antarctic and was initially included in our dataset). The exclusion of taxa with fewer than 55 occurrences represents a methodological choice that strengthens confidence in our findings while highlighting opportunities for refinement. Geographic and temporal biases in GBIF data (
The Arctic region faces an escalating threat from biological invasions driven by increased human activity and climate change. This study employed a data-driven climatic niche horizon scanning approach to identify 2,554 potential new alien vascular plant taxa with climatic niche overlaps in the Arctic under current climate conditions. Six major hotspots were detected, highlighting the pressing need for enhanced biosecurity measures in provinces such as western Alaska, southwestern and southeastern Greenland, northern Iceland, northern Fennoscandia, and Kanin–Pechora. While many potential new alien taxa exhibited low climatic overlap, their establishment could be facilitated by increasing human disturbance, such as in disturbed, nutrient-enriched soils, as well as by climate change impacts. Taxa with high-latitude distributions demonstrated greater potential for occupancy in the Arctic. In a rapidly warming Arctic, some introduced taxa may provide ecological functions or services that support ecosystem resilience, while others may disrupt existing ecological relationships. Management approaches must therefore balance monitoring of potentially harmful invasions with recognition of the inevitability of ecological change in the region. Collaborative efforts among citizens, researchers, policymakers, and stakeholders are essential to develop adaptive management strategies that acknowledge both the risks and potential benefits of newly arriving taxa in the transforming Arctic landscape.
We are grateful to all those who contributed the species occurrence data used in this study and to Brandon Whitley and an anonymous reviewer for their constructive and insightful comments on a previous version of the manuscript.
The authors have declared that no competing interests exist.
No ethical statement was reported.
Grammarly, Microsoft Copilot, and Claude (Anthropic) were used for spell-checking, grammar correction, and suggestions on synonyms to improve readability. During programming, Microsoft Copilot was used to look up functions and packages, while Claude (Anthropic) was used to improve function design and code structure.
This study is part of the BiodivERsA project ASICS (Assessing and mitigating the effects of climate change and biological invasions on the spatial redistribution of biodiversity in cold environments), co-funded by the Research Council of Norway (grant 323304 to KBW) and the Nordic Borealization Network (NordBorN, funded by NordForsk grant number 164079 to JDMS and KBW).
Tor Henrik Ulsted: Conceptualization (supporting); Data curation (lead); Formal analysis (lead); Methodology (equal); Software (lead); Validation (equal); Visualization (lead); Writing – original draft (lead); Writing – review and editing (equal). Kristine Bakke Westergaard: Conceptualization (supporting); Data curation (supporting); Methodology (equal); Funding acquisition (equal); Supervision (equal); Validation (equal); Writing – review and editing (lead). Wayne Dawson: Validation (equal); Writing – review and editing (equal). James D. M. Speed: Conceptualization (lead); Formal analysis (supporting); Methodology (equal); Funding acquisition (equal); Supervision (equal); Validation (equal); Writing – review and editing (equal).
Tor Henrik Ulsted https://orcid.org/0000-0001-8854-2696
Kristine Bakke Westergaard https://orcid.org/0000-0003-4609-8704
Wayne Dawson https://orcid.org/0000-0003-3402-0774
James D. M. Speed https://orcid.org/0000-0002-0633-5595
The dataset and source code used in this study are available as follows:
A full description of the bioinformatics pipeline alongside the source code is available on GitHub (https://doi.org/10.5281/zenodo.15802292).
Raw output files from the climate niche space analysis. The data encompass the complete analytical workflow from initial data wrangling through final post-analysis outputs, providing full transparency and reproducibility for the research (https://doi.org/10.6084/m9.figshare.29402192.v1).
The derived dataset for potential new alien taxa includes GBIF occurrence data for taxa with potential for climatic overlap with the Arctic (https://doi.org/10.15468/DD.2KMYM8).
The derived validation dataset includes GBIF occurrence data for taxa currently found in the Arctic, serving as the validation dataset for this project (https://doi.org/10.15468/DD.Z5B8X5).
Horizon scanning of potential new alien vascular plant species and their climatic niche space across the Arctic
Data type: pdf
CSV datasets
Data type: zip
Explanation note: Zip file with archive containing three CSV datasets with climate niche analysis results and origin–destination connections for vascular plant species in Arctic floristic provinces: (1) included_species.csv—climate analysis results for species included in hypervolume analysis; (2) excluded_species.csv—data for species excluded from final analysis; (3) province_origin_connections.csv—unique taxa counts by origin country and floristic province. See the README file inside the archive for detailed column descriptions and examples.