Diversity and Distribution Patterns of Invasive Alien Plant Species along Dispersal Corridors in Parsa National Park, Central Nepal

Protected areas are vital for preserving native biodiversity, yet invasive alien plant species (IAPS) pose significant threats to their conservation. This study investigates how IAPS diversity varies across their dispersal corridors, such as road verges, riversides, seasonal springs, fire lines, and trekking trails, and examines the relationship between tree canopy openness and IAPS richness. We sampled 156 plots (2m 5m) along these corridors, ensuring a minimum distance of 100 meters between plots. Out of 10 IAPS recorded during the study, road verges and riversides were invaded by 70% of the recorded species, while the walking trail had the lowest number of IAPS (30%). The most prevalent IAPS was Chromolaena odorata, followed by Mikania micrantha and Ageratum conyzoides. One-way ANOVA indicated significant differences in IAPS richness among dispersal corridors (p<0.001). Furthermore, IAPS richness increased with increasing tree canopy openness (R=0.8, p<0.001). Given that road verges and riversides appeared to be the major dispersal corridors of IAPS, it is imperative to prioritize these corridors for IAPS monitoring, early detection, eradication, and control. Such efforts can reduce the establishment probability of new IAPS and mitigate the impacts of the widespread IAPS on the native species and ecosystems. Diversity and Distribution Patterns of Invasive Alien Plant Species along Dispersal Corridors in Parsa National Park, Central Nepal Shreehari Bhattarai 1 *, Balram Bhatta 1, Jeetendra Gautam 1, and Bharat Babu Shrestha 2 1 Faculty of Forestry, Agriculture and Forestry University, Hetauda, Nepal 2 Central Department of Botany, Tribhuvan University, Kathmandu, Nepal *Email: [email protected]

Protected areas are vital for preserving native biodiversity, yet invasive alien plant species (IAPS) pose significant threats to their conservation. This study investigates how IAPS diversity varies across their dispersal corridors, such as road verges, riversides, seasonal springs, fire lines, and trekking trails, and examines the relationship between tree canopy openness and IAPS richness. We sampled 156 plots (2m × 5m) along these corridors, ensuring a minimum distance of 100 meters between plots. Out of 10 IAPS recorded during the study, road verges and riversides were invaded by 70% of the recorded species, while the walking trail had the lowest number of IAPS (30%). The most prevalent IAPS was Chromolaena odorata, followed by Mikania micrantha and Ageratum conyzoides . One-way ANOVA indicated significant differences in IAPS richness among dispersal corridors (p<0.001). Furthermore, IAPS richness increased with increasing tree canopy openness (R=0.8, p<0.001). Given that road verges and riversides appeared to be the major dispersal corridors of IAPS, it is imperative to prioritize these corridors for IAPS monitoring, early detection, eradication, and control. Such efforts can reduce the establishment probability of new IAPS and mitigate the impacts of the widespread IAPS on the native species and ecosystems.

Chromolaena odorata, Ecotourism, Invasive alien species, Protected Area, Tropical forest

Protected areas, such as national parks, are designed to protect native biodiversity and ecosystems while also providing social and economic benefits to local communities and visitors. These areas have multiple objectives, including biodiversity conservation, the protection of cultural heritages, scientific research, recreation, and ecotourism (Esfehani and Albrecht, 2018). Many protected areas also offer important cultural ecosystem services, such as educational opportunities, spiritual enrichment, and recreational activities like hiking and wildlife watching. Protected areas are vital for maintaining natural landscapes and the ecosystem’s integrity, playing a key role in safeguarding species and their habitats (Braun et al., 2016; Chape et al., 2005; Tittensor et al., 2014). However, many protected areas are under threats of multiple factors including the spread of Invasive Alien Plant Species (IAPS) (Foxcroft et al., 2011). The IAPS can have detrimental impacts, affecting native biodiversity, ecosystem health, people’s livelihoods, agriculture, and aquaculture, while also causing economic damage (Aerts et al., 2017; Hattab et al., 2017; Rai and Singh, 2020; Ricotta et al., 2010). In some protected areas, ecotourism, which is often considered as a sustainable means of financing conservation efforts, can unintentionally facilitate IAPS introduction and spread by introducing potentially invasive species into otherwise pristine environments (Tu and Robison, 2013). The encroachment of IAPS into protected areas is an emergent global concern and is increasingly recognized by conservation professionals as a serious and growing threat (Foxcroft et al., 2017; Padmanaba et al., 2017). Invasive alien species are widely regarded as one of the most serious threats to natural ecosystems worldwide, ranking just behind habitat loss and degradation in terms of their impact on native biodiversity (Bellard et al., 2016). Their rapid spread, high reproductive capacity, adaptability to new environments, and competitive dominance over native species have led to habitat degradation and ecosystem disruption. In Nepal, the spread of invasive species such as Chromolaena odorata, Lantana camara, Mesosphaerum suaveolens, Mikania micrantha, and Parthenium hysterophorus is becoming an increasing concern, especially within lowland protected areas (Murphy et al., 2013; Sapkota, 2007; Shrestha et al., 2025). To date, 182 naturalized plant species belonging to 49 families and 129 genera have been recorded in Nepal (Shrestha, 2019; Shrestha and Shrestha, 2021), many of which pose potential ecological threats to the protected areas of Nepal (Bhatta et al., 2020; Bhatta et al., 2024; Shrestha et al., 2025). In particular, the Parsa National Park has already been invaded by more than two dozen of the IAPS, which have reduced regeneration of native trees, altered species composition of plant communities, and modified soil chemistry (Chaudhary et al., 2020). A recent study has also revealed that the IAPS in the Parsa NP are affecting habitat utilization by wild ungulates (Rawal et al., 2025). Within the landscape, including protected areas, dispersal of IAPS propagules often occurs along human-made linear infrastructure such as roads and trails, and natural corridors such as springs and rivers (Brundu et al., 2025; Shrestha et al., 2018), though the protected area boundary may buffer, to some extent, the spread of IAPS to the core areas (Bhatta et al., 2024). Micro-habitats along these linear structures also provide a conducive environment for the establishment of new IAPS, from where they subsequently spread into the landscape (Adhikari et al., 2024). In particular, roads serve as ideal environments and pathways for the spread of IAPS, playing a key role in their establishment and distribution (Le et al., 2018; Meunier and Lavoie, 2012). Despite several research and conservation works focusing on protected areas of Nepal, there is limited information on how linear infrastructure, particularly roads and trails, influences the distribution of IAPS in protected areas of Nepal. In this context, we aim to 1) examine how diversity of IAPS varies across human-made and natural dispersal corridor types in Parsa National Park, and 2) understand how tree canopy openness affects IAPS diversity across potential dispersal corridors. These data can help implement site- and species-specific approaches for the management of IAPS in Parsa National Park and other similar protected areas. 2. Materials and Methods 2.1. Study Area Geographically, Parsa National Park (PNP), located in the south-central lowland Tarai region, ranges from 27º15’ to 27º33’N latitude to 84º41’ to 84º58’E longitude of Nepal extending elevational gradient from 100 to 900 m (Figure 1). Initially established as a wildlife reserve in 1984 to protect the habitat of wild Asian elephants, it was designated as a national park in 2017, covering an area of 627.39 sq. km. To the west of PNP, there is another protected area the Chitwan National Park which is also a World Heritage site. The dominant tree species in the park is Sal ( Shorea robusta ). The soil in the area is primarily gravelly and highly porous, characterized by a low water table, with many streams disappearing into the permeable sediments. Despite this, the park supports a rich biodiversity, including 298 species of vascular plants, 37 mammals, 503 birds, 8 species of herpetofauna, and 8 species of fish (Bhuju et al., 2007). The Parsa-Chitwan Complex spans the Himalayan foothills and includes floodplains, Dun valleys, the Bhabar tract, and the Siwalik and Chure hills. Figure 1. Map of Parsa National Park showing study sites The study area is predominantly covered (approximately 90%) by tropical and subtropical forests, primarily dominated by Sal ( Shorea robusta ) and its associated species. However, vegetation composition varies noticeably with changes in slope, elevation, soil type, and microclimatic conditions. Several forest types are present within the landscape, such as mixed deciduous riverine forest, characterized by species such as Sissoo ( Dalbergia sissoo ), Khair ( Senegalia catechu ), Simal ( Bombax ceiba ), Phorsa ( Grewia disperma ), and Khirro ( Holarrhena pubescens ), accompanied by shrub species like Bayer ( Zyzyphus mauritiana ), Boksi Kanda ( Caesalpinia acucullata ), and Asare ( Murraya koenigii ). Similarly, a mixed deciduous hardwood forest comprising Sal ( Shorea robusta ), Haldu ( Adina cordifolia ), Saj ( Terminalia alata ), Tantari ( Dillenia pentagyna ), Botdhairo ( Lagerstroemia parviflora ), Sindure ( Mallotus philippensis ), and Kurilo ( Asparagus racemosus ), along with climbers such as Ram Datiwan ( Smilax ovalifolia ) and juvenile Phoenix species. 2.2. Data collection A reconnaissance survey was conducted together with the staff of PNP for the identification of different possible trails within the core area of PNP, where transect lines were set along the trails of Bhata-Laukidaha, Bhata-Amlekhgunj fire line, Rambhouri-Ghodemasan, Bhata-Rambhouri, Bhata Range post- Dharmasala, Bhata-Devakidaha, Bhata-Jamini river, Bhata-Chabi river and Grassland areas. Altogether 156 plots of size 2m*5m were sampled at intervals of >100 m along different dispersal corridors such as permanent riverside (N=34), seasonal spring (N=32), fire line (N=30), graveled road (N=27), and walking trail (N=33). The tree canopy in each plot was measured using a densiometer. 2.3. Data analysis The frequency and abundance of each species were calculated by using the methods described in (Dallmeier, 1992; Shukla and Chandel, 2000). \begin{equation} \text{Frequency\ }\left(F\right)=\frac{\text{Total\ number\ of\ quadrats\ in\ which\ the\ species\ occur}}{\text{Total\ number\ of\ quadrats\ studied}}\times 100\nonumber \\ \end{equation}\begin{equation} \text{Abundance\ }\left(A\right)=\frac{\text{Total\ number\ of\ individuals\ of\ a\ species\ in\ all\ the\ quadrats}}{\text{Total\ number\ of\ quadrats\ in\ which\ the\ species\ occurs}}\nonumber \\ \end{equation} The one-way ANOVA was used to compare the IAPS richness among different trail types. The influence of the canopy openness on the IAPS richness was assessed using regression analysis. To assess patterns in IAPS community composition across different trail types, a Non-metric Multidimensional Scaling (NMDS) analysis was performed (Bray and Curtis, 1957; Kuswantoro et al., 2020). The Bray-Curtis dissimilarity index was used to construct the distance matrix. NMDS ordination was then carried out using a two-dimensional solution to provide a simplified yet accurate representation of multivariate relationships among sites. Ordination stress values were examined to evaluate the adequacy of the two-dimensional configuration, with stress values close to or less than 0.2 considered to indicate an excellent representation of the underlying data structure (Clarke, 1993; Dhakal et al., 2024). Site scores were overlaid with trail type groupings to visualize clustering patterns in species composition among Fire line, Roadside, River side, Walking trail, and Seasonal spring categories. The analysis was performed in R (version 2025.05.1) using the vegan package. 3. Results 3.1. IAPS richness along different trails The one-way ANOVA revealed significant variation in invasive alien plant species (IAPS) richness among different trail types (F = 15.70, p < 0.001). Tukey’s post-hoc comparisons showed that roadside habitats supported significantly higher IAPS richness compared to permanent riversides, seasonal springs, and walking tracks, while the walking tracks harbored the lowest richness, being significantly different from all other trail types (Figure 2). Permanent riverside, seasonal spring, and fire line trails exhibited intermediate and statistically comparable levels of IAPS richness. Figure 2. IAPS richness along different trail types The Non-metric Multidimensional Scaling (NMDS) analysis of invasive species abundance across different trail types revealed clear patterns in community composition (Figure 3). The NMDS ordination achieved a stress value of 0.059, indicating an excellent fit of the two-dimensional representation to the original multidimensional data. Sites belonging to similar trail types tended to cluster together, suggesting comparable species composition within each trail category. For example, Fire line and Roadside sites clustered closely, driven primarily by high abundances of Chromolaena odorata, Mikania micrantha, and Ageratum conyzoides . River-side sites formed a distinct cluster, reflecting differences in dominant species composition relative to other trail types. Walking trail and Seasonal spring sites were more dispersed, indicating greater variability in species abundance. Figure 3. NMDS of invasive alien plant species abundance across trail types 3.2. Frequency and Abundance of IAPS along different trails Among the IAPS, Chromolaena odorata shows the highest frequency, with 100% occurrence along roads and 87.5% along permanent riversides (Table 1). Mikania micrantha is also prevalent, particularly along roads and riversides whereas Ageratum conyzoides colonized frequently along roads and riversides. In contrast, species like Spermacoce alata and Senna tora have minimal occurrences, mostly confined to seasonal springs with a low frequency of 2.78%. Table 1. Frequency of IAPS along different trails | 1 | Chromolaena odorata (L.) R. M. King & H. Rob. | 87.50 | 80.56 | 56.25 | 100.00 | 8.70 | | 2 | Spermacoce alata Aubl. | 0.00 | 2.78 | 0.00 | 0.00 | 0.00 | | 3 | Senna tora (L.) Roxb. | 0.00 | 2.78 | 0.00 | 0.00 | 0.00 | | 4 | Mikania micrantha Kunth | 66.67 | 41.67 | 28.13 | 100.00 | 13.04 | | 5 | Ageratum conyzoides L. | 54.17 | 16.67 | 34.38 | 87.50 | 8.70 | | 6 | Ageratum houstonianum Mill. | 0.00 | 0.00 | 0.00 | 25.00 | 0.00 | | 7 | Mimosa pudica L. | 0.00 | 0.00 | 3.13 | 25.00 | 0.00 | | 8 | Parthenium hysterophorus L. | 0.00 | 0.00 | 0.00 | 25.00 | 0.00 | | 9 | Amaranthus spinosus L. | 0.00 | 0.00 | 0.00 | 12.50 | 0.00 | | 10 | Senna occidentalis (L.) Link | 4.17 | 25.00 | 6.25 | 0.00 | 0.00 | Ageratum conyzoides stands out with the highest abundance, particularly along walking tracks (22.00) and fire lines (17.55) (Table 2). Chromolaena odorata also shows high abundance, especially along permanent riversides and fire lines. Mikania micrantha is relatively abundant across all trails, with values ranging from 7.13 along roads to 12.69 along permanent riversides. In contrast, species like Spermacoce alata, Senna tora, and Mimosa pudica have very low abundance, being mostly confined to temporary riversides or specific areas with minimal presence. Table 2. Abundance of IAPS along different trails | 1 | Chromolaena odorata (L.) R. M. King & H. Rob. | 14.38 | 10.07 | 14.33 | 13.50 | 9.50 | | 2 | Spermacoce alata Aubl. | 0.00 | 2.00 | 0.00 | 0.00 | 0.00 | | 3 | Senna tora (L.) Roxb. | 0.00 | 3.00 | 0.00 | 0.00 | 0.00 | | 4 | Mikania micrantha Kunth | 12.69 | 8.93 | 9.44 | 7.13 | 10.33 | | 5 | Ageratum conyzoides L. | 18.92 | 13.00 | 17.55 | 17.29 | 22.00 | | 6 | Ageratum houstonianum Mill. | 0.00 | 0.00 | 0.00 | 17.00 | 0.00 | | 7 | Mimosa pudica L. | 0.00 | 0.00 | 2.00 | 3.50 | 0.00 | | 8 | Parthenium hysterophorus L. | 0.00 | 0.00 | 0.00 | 3.50 | 0.00 | | 9 | Amaranthus spinosus L. | 0.00 | 0.00 | 0.00 | 7.00 | 0.00 | | 10 | Senna occidentalis (L.) Link | 2.00 | 5.67 | 6.00 | 0.00 | 0.00 | 3.3. Effects of the canopy on the IAPS richness The influence of canopy openness (%) on the IAPS richness revealed a strong and statistically significant (R=0.8, p=0.000) role. At 100% canopy openness, there was a notable increase in IAPS diversity (Figure 4), indicating that fully open canopies support higher levels of invasive species diversity. Conversely, as canopy openness decreased, IAPS diversity also declined significantly, with many areas showing no invasive species under very dense canopy conditions. Figure 4. Influence of canopy openness to invasive alien plant species richness 4. Discussion 4.1. Diversity and distribution pattern of IAPS along trail types The findings reveal a distinct pattern in the distribution of IAPS, with roadsides and permanent riversides identified as hotspots for both IAPS richness and abundance. The high proportion of alien species found along roads (70%) and permanent riversides (60%) highlights these environments as significant pathways for the spread of invasive species. Roads and rivers are typically associated with disturbances that promote the establishment of invasive species, including habitat fragmentation, altered microclimatic conditions, and increased human activity which often serve as corridors for dispersal and provide favorable habitats for exotic species (Parendes et al., 2000; Silva et al., 2017). The presence of roads and rivers can contribute to the invasion of protected areas in two key ways: by facilitating the migration of alien plants through external human activities, such as transportation by people and vehicles (Niggemann et al., 2009; Von der Lippe and Kowarik, 2007), and by creating novel environments conducive to the establishment of alien species (Greenberg et al., 1997). These results suggest that disturbance intensity and type strongly influence invasion patterns, with roadsides acting as major invasion hotspots, likely due to continuous anthropogenic disturbance and propagule pressure, whereas walking tracks are less susceptible to invasion. The widespread presence of Chromolaena odorata across all trails highlights its adaptability to highly disturbed environments, where it can outcompete native vegetation. Similarly, Mikania micrantha also demonstrates a strong presence, showcasing its ability to thrive in wet and disturbed conditions. The prevalence of these species in disturbed areas like roadsides and riversides indicates that such habitats offer ideal conditions for their rapid invasion and colonization. Their dominance poses significant ecological challenges, including the alteration of native plant communities, reduction in biodiversity, and interference with the regeneration of native species (Banerjee and Dewanji, 2017). In contrast, walking tracks, which have only 30% IAPS and the lowest diversity index (0.053), reflect lower disturbance levels. These tracks, being narrower and less frequently disturbed compared to roads, provide fewer opportunities for the introduction and colonization of invasive species. This suggests that less-disturbed habitats, such as walking tracks, maintain higher resistance to invasion, thereby preserving native species and sustaining ecological balance. In contrast, species like Spermacoce alata and Senna tora have minimal occurrences, mostly confined to seasonal springs with a low frequency of 2.78%. Other species, such as Ageratum houstonianum, Mimosa pudica, and Parthenium hysterophorus are limited to specific areas, particularly along roads with moderate to low frequencies. This pattern reveals a concentration of invasive species in highly disturbed areas like roads and riversides, while other trails like walking tracks exhibit much lower invasion rates. Ageratum conyzoides stands out with the highest abundance, particularly along walking tracks and fire lines, suggesting that species such as Ageratum conyzoides and Chromolaena odorata dominate the landscape, while other species exhibit more restricted distributions based on trail types. Species labels on the NMDS plot highlight the key species contributing to community differences, whereas species absent from all sites were automatically excluded from the analysis. Overall, the NMDS results indicate that trail type strongly influences the composition of invasive plant communities, with certain species acting as indicators of specific disturbance regimes. The ellipses around each trail type further emphasize the clustering patterns, showing that sites within the same trail type share similar species assemblages. 4.2. Relationship between canopy openness and IAPS richness The strong positive correlation (R=0.8, p=0.000) between canopy openness and the diversity of IAPS highlights the crucial role of light availability in fostering the spread of these species. At 100% canopy openness, where light is plentiful, IAPS diversity reaches its peak, indicating that fully open environments are particularly vulnerable to invasion. Invasive species, which are often fast-growing and require ample light, can establish and proliferate rapidly in open-canopy areas where competition from native plants is diminished and environmental conditions such as light, temperature, and moisture are more conducive to their growth (Le et al., 2018). Forest canopy cover is the overriding biotic covariate suppressing the diversity and distribution of IAPS (Sharma et al., 2022), and an increase in canopy cover and closure of forest gaps through proper management of degraded forests can prevent plant invasions and suppress the growth of previously established IAPS (Khaniya and Shrestha, 2020; Subedi et al., 2024). These results suggest that greater canopy openness is linked to higher IAPS diversity, while reduced canopy openness limits the presence and spread of invasive species, likely due to decreased light availability or other ecological constraints. Disturbances that increase canopy openness, such as logging, natural disasters, or land-use changes, heighten the risk of IAPS invasion. Thus, maintaining or restoring forest ecosystems in degraded areas can be an effective strategy to curb the spread of invasive species and protect native biodiversity. 4.3. Implications for conservation Our findings indicate that the distribution and diversity of IAPS were concentrated along roads, suggesting that movement along these routes and the presence of open canopy conditions facilitate the spread and colonization of IAPS. The results underscore the urgency of addressing plant invasions in the PNP, which are becoming a serious issue and necessitate immediate IAPS-targeted management actions. The study highlights that trail runners and vehicles may serve as vectors for seed dispersal and that sporting events within protected areas can increase the risk of alien plant introductions. To mitigate these threats, protected area authorities should implement biosecurity measures and raise awareness in collaboration with ecotourism managers. Given that roads and riversides act as corridors enhancing IAPS invasion, prioritization of these dispersal corridors for monitoring, early detection, eradication, and control of the IAPS and integrating the management of IAPS in the park’s management plan is crucial to prevent the introduction of these harmful species and to conserve native biodiversity. Careful management of IAPS, implementing strategies such as regular monitoring, early detection of IAPS, habitat restoration, and public awareness, is mandatory to minimize the potential risk 5. Conclusions The study underscores the critical role of disturbance regimes and canopy structure in shaping the diversity and distribution of invasive alien plant species (IAPS) within Parsa National Park. Roadsides and riversides emerge as primary invasion hotspots due to their openness and frequent disturbances, while intact forest canopies and minimally disturbed walking tracks exhibit higher resistance to invasion. The strong positive relationship between canopy openness and IAPS richness further highlights the vulnerability of degraded or open habitats to invasion. These findings emphasize the urgent need for integrated management approaches, including the restoration of forest canopy, targeted monitoring along dispersal corridors, and the implementation of strict biosecurity and awareness programs. Such efforts are essential to mitigate the spread of IAPS, safeguard native biodiversity, and maintain the ecological integrity of protected areas like PNP. Author contributions SB: conceptualization, data curation, formal analysis, methodology, writing - original draft, writing- review and editing. BB: conceptualization, investigation, methodology, supervision, writing - review and editing. JG: conceptualization, investigation, methodology, writing - review and editing. BBS: conceptualization, investigation, methodology, supervision, validation, writing - review and editing. Acknowledgments The authors are thankful to Prakash Pun, Minu Gautam, Yadav Shahi Thakuri, Chandan Sah, PNP officials, and local people of the buffer zone for their cooperation and continuous help in the fieldwork. Equally, they are grateful to the Department of National Parks and Wildlife Conservation, Nepal, for providing permission to study in the park, and Faculty of Forestry (AFU) and the Directorate of Research and Extension (AFU) for providing necessary support during this study. Conflicts of interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper. Data availability statement All the required data are uploaded as supplementary materials.

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Authors Metrics & Citations Metrics Article Usage 415views 179downloads Citations Download citation Shreehari Bhattarai, Balram Bhatta, Jeetendra Gautam, et al. Diversity and Distribution Patterns of Invasive Alien Plant Species along Dispersal Corridors in Parsa National Park, Central Nepal. Authorea. 19 November 2025. DOI: https://doi.org/10.22541/au.176357021.10290091/v1 DOI: https://doi.org/10.22541/au.176357021.10290091/v1 If you have the appropriate software installed, you can download article citation data to the citation manager of your choice. Simply select your manager software from the list below and click Download. For more information or tips please see 'Downloading to a citation manager' in the Help menu.