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10 September 2026

Trait Plasticity and Habitat–Environment Heterogeneity Explain the Landscape-Scale Invasion Success of Verbesina encelioides (Cav.) Benth. & Hook. f. Ex A. Gray and Its Implications for Ecosystem and Biodiversity Management in the Hindu Kush Region

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and
1
Department of Botany, University of Malakand, Chakdara P.O. Box 18800, Pakistan
2
School of Applied Sciences, University of the West of England, Frenchay Campus, Coldharbour Lane, Bristol BS16 1QY, UK
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Authors to whom correspondence should be addressed.
Sustainability2026, 18(18), 9305;https://doi.org/10.3390/su18189305
This article belongs to the Special Issue Plant Ecological Function Research and Ecological Conservation
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Abstract

Functional trait plasticity enables invasive plant species to establish and persist across diverse environmental conditions. Following the extensive habitat disturbance caused by the 2010 floods in Khyber Pakhtunkhwa, Verbesina encelioides rapidly spread into disturbed and natural habitats. This study evaluated the phenotypic plasticity and biomass allocation traits of Verbesina encelioides across five contrasting habitats (i.e., cropland, roadside, riverside, urban, and abandoned land). Habitat differences were assessed using analysis of variance with Tukey’s HSD post hoc test, while multivariate analysis was used to examine relationships among functional traits, environmental variables, and habitats. Log-transformed data were used for linear regression analyses to satisfy normality assumptions. Plant functional traits varied significantly among habitats, demonstrating substantial phenotypic plasticity. Riverside vegetation had high functional trait plasticity followed by abandoned habitats. In contrast, riverside populations had fewer branches (14.3 ± 1.4), leaves (184.7 ± 10), lower leaf area (5360 ± 77 cm2), and reduced inflorescence biomass (8.61 ± 2.1 g) than abandoned-land populations. Cropland plants were taller (106 ± 3 cm) with heavier seeds (0.30 ± 0.04 g; p < 0.05), whereas roadside plants had smaller leaves (6.1 ± 0.3 cm) but produced more flowers (38.4 ± 6.2; p < 0.001). Biomass allocation remained relatively stable across habitats (p > 0.05), with greater investment in aboveground than belowground structures. Soil texture, organic matter, temperature, and precipitation were the principal environmental variables associated with trait variation, indicating that local environmental conditions strongly influence plant performance (p < 0.05). PCA revealed that seed traits dominated the first axis and were associated with cropland and riversides. Redundancy analysis indicated that soil properties (sand, silt, pH, EC) and climatic factors (precipitation, temperature) significantly influenced trait distribution. Hierarchical cluster analysis shows that riverside was distinct from other clusters both in functional traits and environmental variables. Inter-trait correlation revealed a significant relationship between leaf and seed traits. Overall, these findings reveal that V. encelioides adjusts its growth and reproductive traits according to habitat conditions. Greater seed weight in croplands may enhance seedling establishment under competitive conditions, whereas increased flower production in disturbed roadside habitats may promote dispersal and reproductive success of the plant. The strong functional trait plasticity of V. encelioides across different habitats highlights its potential to persist under habitat disturbance and changing environmental conditions, therefore posing risk to native plant communities and ecosystem structure. These findings support the need for habitat-based monitoring, protection of native flora and restoration of disturbed areas as part of sustainable land and biodiversity management.

1. Introduction

Anthropogenic climate change has caused an increase in global average temperatures and changed precipitation patterns over the past century due to the accumulation of greenhouse gases from human activities [1]. These climate changes profoundly impact plant phenology, including abundance, distribution, and community composition [2], thereby altering environmental conditions and creating novel ecological niches. Changes in temperature and precipitation regimes, together with increasing environmental disturbances, can change habitat suitability and resource availability, and competitive interactions within plant communities facilitate the establishment, persistence, and spread of invasive plant species [3,4]. Consequently, climate change may not only alter the distribution of invasive plants but also enhance their impacts on ecosystem structure and function, thereby threatening biodiversity, human health, and economic development [5]. Phenotypic plasticity is the capability of species to produce distinct phenotypes in response to varying environmental conditions [6]. Plant species with high phenotypic plasticity are able to opportunistically invade heterogeneous environments and expand their realized niche [7,8]. Plant species deemed “invasive” often exhibit high plasticity in vegetative and reproductive biomass, thereby enhancing their colonization ability and fitness under diverse environmental conditions [9,10]. Previous studies have shown that invasive plant species often possess higher growth rates, prolonged flowering periods, greater fecundity, efficient seed dispersal, and enhanced reproductive output than co-occurring native species, making them successful invaders which prove difficult to control in new regions [11]. A recent assessment recorded 392 invasive vascular plant species across South Asia, where Lantana camara and Ageratina adenophora have been reported to reduce native understory herb and shrub species richness by 29–40% in Chir pine forests of the central Himalayas [12]. Similarly, invasion success of the four invasive species [13] including Xanthium [14] and some weeds [15] has been reported in sub-mountain valleys and plains of Khyber Pakhtunkhwa, which is an important concern for ecosystem health and biodiversity conservation. Understanding the response of invasive species to changing environmental conditions can help us to monitor and protect vulnerable plant communities.
The functional traits of invasive plants vary depending on the habitat type in which they grow [16]. Among these traits, the shoot and root attributes are useful morphological indicators for successful invasion of plants across diverse habitats [17]. Along with morphological traits, physiological traits such as photosynthetic capacity also play an important role in the growth and competitive performance of invasive plants [18]. Moreover, several studies have found that invasive species exhibit higher specific leaf area and leaf dry matter content in certain habitats [19], which were considered to be the key predictors of the plant’s overall strategy, particularly its capacity for rapid resource acquisition and invasion success [20]. Reproductive traits also exhibit considerable plasticity, allowing invasive species to maximize seed production and population growth under favorable habitat conditions [21]. Habitats with greater resource availability tend to be more productive for invaders, leading to greater reproductive output and faster dispersal [22]. Moreover, invasive plants can adjust carbon allocation among roots, shoots, and reproductive structures according to resource availability and environmental stress. Root–shoot and source–sink dynamics regulate the distribution of assimilated carbon with greater investment in roots under water or nutrient limitation and increased allocation to aboveground growth and reproduction when resources are more favorable, increasing the survival and reproductive success across contrasting habitats [23]. However, many invasive species expand to managed or disturbed areas, particularly agricultural fields and roadsides [24,25]. Some of these invasive species subsequently spread into less disturbed natural habitats, which adversely affects ecosystem processes and functions [26,27]. Such habitat disturbance can increase invasion risk and create additional challenges for sustainable management and natural ecosystems. Identifying habitats vulnerable to invasion can help us to focus our monitoring, management, and restoration efforts. Given the ongoing expansion of invasive plant species, detailed research is required to explore their morphological and reproductive traits and their responses to habitat–environment heterogeneity to explain their invasion patterns across interconnected landscapes.
Among the rapidly expanding invasive species, Verbesina encelioides (Cav.) Benth. & Hook. f. ex A. Gray (Asteraceae), commonly known as golden crown-beard or wild sunflower, has attracted increasing attention because of its remarkable ecological adaptability and invasion success. The species is native to the southwestern United States and the Mexican Plateau but has now spread rapidly across several continents, including Asia, Africa, Europe, and India [28,29]. It has become naturalized in many tropical and subtropical regions, where it invades roadsides, railway tracks, abandoned agricultural lands, wastelands, urban areas, field crops, and other disturbed habitats [30]. Its invasion has been reported in the semi-arid region of India [31], and cropland and roadside areas in Morocco [32]. The successful establishment of the species across contrasting habitats is linked to its high phenotypic plasticity, reproductive ability, and effective dispersal and persistence of seeds. A mature V. encelioides plant typically produces 29–254 capitula, each containing approximately 300–350 seeds, resulting in several thousand seeds produced per plant during a single growing season [33], supporting its invasion success. The species also exhibits prolonged seed viability, with seeds remaining viable in the soil for up to four years [34]. Seed dispersal occurs through wind, water, animals, and vehicular movement, and due to their light weight and winged margins, seeds can disperse over long distances. Some seeds, by contrast, remain buried beneath the parent plant and germinate during subsequent favorable seasons, ensuring continuous recruitment and long-term persistence within invaded habitats [35]. In addition, the species exhibits considerable phenotypic plasticity, enabling it to flower and produce viable seeds even when plants are only 20 to 25 cm tall under field conditions [36,37]. Such reproductive flexibility allows populations to persist even under environmental stress and frequent disturbance. In Pakistan, V. encelioides has emerged as an increasingly problematic invasive weed during the past decade. Field observations indicate extensive colonization in Khyber Pakhtunkhwa after the 2010 devastating floods. The floodwater may have carried its seeds downstream and deposited sediments (hydrochory), where the newly disturbed and open riverbank areas created favorable sites for seed germination and establishment. However, direct evidence linking the floods to the initial spread of V. encelioides is still limited. The plant densely occupied riverbanks, subsequently expanding into nearby agricultural fields, roadsides, wastelands, grazing lands, urban areas and other disturbed habitats. Despite the rapid expansion of V. encelioides and its increasing ecological and economic importance, the mechanisms of its successful invasion across different habitats remain poorly understood. Previous studies have largely focused on taxonomy, distribution, biology and weed management including manual removal, cultural practices, and herbicide-based control of Verbesina encelioides in invaded areas. However, the habitat-specific functional trait plasticity of V. encelioides and its relationship with environmental heterogeneity remain poorly understood, particularly in Khyber Pakhtunkhwa. Addressing this gap is important for understanding how V. encelioides responds to disturbed and heterogeneous habitats and how its invasion may affect native vegetation. This information can support habitat-based monitoring, sustainable land management, and restoration of degraded areas affected by invasive plants. Therefore, this study examines how functional trait plasticity and habitat–environment heterogeneity contributes to the landscape-scale invasion success of V. encelioides in Khyber Pakhtunkhwa. We hypothesize that variation in environmental conditions across different habitat types drives significant changes in vegetative and reproductive traits and biomass allocation, where plants show greater reproductive investment (flower and seed production) in highly disturbed habitats, while vegetative traits dominate in less disturbed habitats which enable the plant to utilize resources and maintain fitness under diverse ecological conditions. Specifically, the objectives of this research were: (i) to evaluate the variations in both vegetative and reproductive functional traits across different habitat types; (ii) to determine the belowground (root) and aboveground (vegetative and reproductive) dry biomass allocation among habitats; and (iii) to identify key environmental factors that may affect the functional traits and biomass of V. encelioides across different habitats.

2. Materials and Methods

2.1. Study Area

This research was conducted in V. encelioides-dominated areas across Khyber Pakhtunkhwa, Pakistan (Figure 1). The province borders Baluchistan, Punjab, Azad Jammu and Kashmir (AJK) and Afghanistan, spanning from 34.22° to 35.22° N and 71.01° to 72.40° E with elevations ranging from 299 to 1525 m above sea level [38]. The study area is characterized by diverse topography, encompassing fertile plains and rugged mountainous regions that support a wide range of environmental conditions and vegetation. Lower elevation zones include human settlements and agricultural activities, and have an abundance of alien species, while the high-altitude zone is predominantly covered by subtropical pine and deciduous forests [39].
Figure 1. Spatial distribution of sampling points across the five habitat types: riverside, cropland, abandoned land, roadside, and urban areas. Each habitat type is represented by a different color.
Agriculture is the primary contributor to the province’s GDP, which relies mainly on precipitation, with infrequent irrigation [40]. The main crops of the area are wheat, maize, rice, sugarcane, tobacco, barley, gram, sugar beet, millet, and many types of vegetables and fruits [41]. The province experiences considerable climatic variation, with hilly regions having cold winters and cool summers, while the plains are warmer year-round (Figure 2). Thirty-two years of climate records (precipitation, temperature and relative humidity data) obtained from the Pakistan Meteorological Department indicate uniform periodicity with a slight variation in temperature (0.4 °C per year) on the plain [14].
Figure 2. Annual variations in major climatic variables in the study region.

2.2. Field Study

Habitat Types and Environmental Factors

We surveyed the population of V. encelioides in five different habitats—riverside, cropland, abandoned land, roadsides, and urban areas—where the plant was abundantly present (Figure 3 and Figure 4). We randomly selected ten plots (5 m × 5 m = 25 m2) within each habitat when the growing season was at peak (July –September) for later measurements. A 5 × 5 m plot was used because V. encelioides forms patchy, heterogeneous stands, allowing representative phytosociological assessment of the invasive population and associated vegetation. In each invaded site, we collected 3 kg soil samples (2 kg from opposite diagonal sides and 1 kg from the center) and subsequently pooled them using auger borings for soil physiochemical analysis following standard pedological protocols [42,43]. We sampled the soil from 0–30 cm depth, which is typically the nutrient-rich layer where most of the biological activity, root growth, and nutrient cycling take place [44].
Figure 3. V. encelioides infestation across different habitats of Khyber Pakhtunkhwa.
Figure 4. Schematic flow illustrating the overall research methodology.
We measured soil electrical conductivity and pH in situ in a soil–water suspension (1:5) using an EC-meter (Model CC601 Century) and digital pH meter (Model CON.3173) respectively. In the laboratory, soil samples were air-dried and then passed through a 2 mm sieve for textural classification (sand, silt and clay %) following USDA methods. Lime% was obtained using the geometric method [45]. Soil organic matter content was determined using the Walkley–Black method [46], whereas available phosphorus (P2+), total nitrogen (N), and exchangeable potassium (K+) were determined using the micro-Kjeldahl technique [47]. Since the local station meteorological data for each stand were not available, we used gridded climate data (from 1981–2021) derived online from NASA (https://power.larc.nasa.gov/data-access-viewer/) (accessed on 15 august 2024) with a spatial resolution of 0.5° × 0.5° to generate more biologically meaningful variables. The climatic data included represent annual trends (e.g., maximum (Tmax) and minimum (Tmin) temperature, precipitation (mm), and % relative humidity), dew/frost point (DEW), surface soil wetness (SSW), root zone soil wetness (RZSW), and surface pressure (SPr) in kPa. The mean temperature (MT) was calculated as the average annual temperature, while the growing season temperature (GST in °C) and growing season precipitation (GSP in mm) were averaged from March to October [48].
Summer drought severity index (SDSI) was calculated using the following formula followed by [49]:
SDSI = PE/ME
where PE is the sum of rainfall (in mm) during June–August and ME is mean maximum temperatures (in °C) of three months (in June, July, and August).

2.3. Measurements of Functional Traits

Within each selected plot, we randomly selected ten mature and healthy V. encelioides plants at the end of the flowering stage (July_September) for functional trait analysis; i.e., a total of 500 plants were screened (10 plants per plot × 10 plots per habitat × 5 habitats). Characteristics including plant height (PHe), number of branches (NoB), number of leaves (NoL) and number of flowers per plant (NoF) were measured manually during the sampling phase. After these measurements, each of the 10 mature and healthy plants per site was uprooted and brought to the laboratory in polythene bags. Soil attached to the roots was gently removed by hand, and the roots were then carefully washed with low-pressure tap water over a fine-mesh sieve to prevent losing fine roots. After washing, root length and biomass were measured [50]. Reproductive and vegetative traits were evaluated to assess adaptability in plant functional types (PFTs), following [51]. Root length (RL), average leaf length (ALL), and average leaf width (ALW) were measured using scales, while average leaf thickness (ALT) and stem diameter (SD) were measured using digital Vernier calipers. Leaf area index (LAI) and leaf area per plant (LAPP) were calculated according to the following formula [52,53]:
LAI = L × W ×K
where L = length of leaf, W = width of leaf, and K = constant of 0.75. The use of a correction factor (0.75) is consistent with previous approaches applied to members of the family Asteraceae [54].
Leaf area per plant = Mean leaf area × Number of leaves per plant
Specific leaf area (SLA) was measured by dividing leaf area by leaf dry weight [55]. Among the reproductive traits, number of seeds (SN) was measured by counting three randomly selected flowers from each plant and then averaged to get mean values. Seed length (SL), seed width (SWd) and seed diameter (SDi) were measured using a scale and digital Vernier calipers respectively, while seed weight (SW) was determined using a digital balance (Mettler Toledo-ME104, accuracy = 0.0001 g). For biomass allocation, each collected plant per site was separated into functional parts (roots, stems, leaves and flowers) and dried at 60 °C for 72 h [56] and then weighed using a digital balance. Dry matter content (g) was used to calculate biomass allocation following [57].

2.4. Statistical Analysis

All data were analyzed and graphically represented using OriginPro software (Version 2024, OriginLab Corporation, Northampton, MA, USA). Analysis of descriptive statistics and analysis of variance were performed in OriginPro softwere (Version 2024) to determine the inter-habitat-level variation in plant functional traits and environmental variables. Difference in group mean was determined using Tukey’s honest significance test with significance level set at p ≤ 0.05. Grubbs’ test was applied to detect outliers in plant functional trait data. A few outliers were found in cropland and roadside root traits but were retained in the subsequent analysis due to natural biological variation. The functional trait data across each habitat were subjected to principal component analysis in OriginPro software (Version 2024) following a previously adopted approach [58]. In the PCA biplot, the habitats are plotted as observations while functional traits are displayed as loading vectors. Redundancy analysis was conducted using PC-ORD software (Ver. 6.0) to evaluate the relationships among habitat-level soil variables and functional traits.
Additionally, hierarchical cluster analysis (HCA) and heatmap analysis were performed in Origin Pro 2024 to assess habitat similarity based on soil variables and V. encelioides functional traits using Pearson correlation distance and the UPGMA clustering method. Before conducting inter-trait analysis, the normality distribution of functional trait data was evaluated using the Shapiro–Wilk test at significance level α = 0.05. Most functional traits exhibited significant deviation from normality assumptions (p < 0.05) (Table S1); therefore, we log-transformed (log10(x)) all the functional trait data to fit a normal distribution before further analysis. The relationships among functional traits were analyzed using the allometric linear model of regression in GraphPad Prism (Ver 9.0).

3. Results

3.1. Functional Trait Variations Among the Habitats of V. encelioides

The descriptive statistics of both vegetative and reproductive traits of V. encelioides are provided in Table 1. Several vegetative traits differed significantly among habitats (p < 0.01). Riverside plants had the greatest stem diameter (9.80 ± 0.77) and leaf area index (29.09 ± 4.11), whereas plants growing on abandoned land had the highest numbers of branches (46.31 ± 9.30) and leaves (288.27 ± 43.98), as well as the greatest leaf area per plant (6116.8 ± 653.6). Likewise, leaf length and leaf width show significant variation (p < 0.05) among the habitats with shorter ALL (6.15 ± 0.3) and narrow ALW (3.25 ± 0.2) found on roadside plants. The height of V. encelioides shows significant differences, with higher values in the cropland habitat (106 ± 3.9) and lower values (79.2 ± 5.19) in abandoned land. The other vegetative traits and VDBM do not differ significantly between habitats.
Table 1. Descriptive statistics of functional traits of V. encelioides across different habitats in Khyber Pakhtunkhwa. One-way analysis of variance followed by Tukey’s HSD post hoc test was used for significant differences among the habitat types.
Reproductive output and seed characteristics were strongly influenced by habitat heterogeneity. Number of flowers varied significantly among habitats (p < 0.01), with the highest production at roadsides (38.41 ± 6.28) and the lowest in croplands (12.91 ± 2.08). Seed number and seed weight also showed significant differences among habitats (p < 0.05) with the highest seed number recorded in riverside and cropland habitats (143.52 ± 9.43 and 140.87 ± 11, respectively), while seed weight was highest in cropland habitats (Table 1). In contrast, abandoned land showed low seed number and weight, whereas other traits like seed dimension (SL, SWd, SDi) and inflorescence dry weight did not exhibit significant variations among the habitats.
The variations in dry biomass among the habitats are shown in Figure 5. Regardless of habitat type, V. encelioides allocated more biomass to aboveground (vegetative and reproductive structures) tissues compared to belowground organs. Biomass allocation to roots was higher at roadside, followed by riverside and abandoned land. However, the biomass allocations to vegetative, reproductive and total plant were slightly greater in abandoned fields, followed by riverside and roadside, while lower biomass allocation was recorded in cropland. Overall, no significant differences were observed among the habitats for any of the measured biomass.
Figure 5. Biomass allocation of Verbesina encelioides based on dry matter (g) across different habitats. Values are means (n = 10) and the bars represent the standard deviation. Within each parameter, values with the same letter are not significantly different at 0.05 level using Tukey’s HSD test. Note: BDB (belowground dry biomass), VDB (vegetative dry biomass), RDB (reproductive dry biomass), TPDB (total plant dry biomass).

3.2. Influence of Environmental Variables Across Habitats

The environmental variables, including topographic, edaphic, physicochemical properties and climatic factors, are presented in Table 2. Environmental variables like elevation (F = 3.19; p = 0.02) and latitude (F = 4.80; p = 0.003) differed significantly between the habitats (Figure 6a,b). Croplands were at higher elevation while urban areas were at lower elevations. The soil of the cropland habitat contained the highest percentage of silt (69.40%) and organic matter (2.74%) In contrast, the highest sand content (F = 57.18; p < 0.0001) was found in roadside soil (69.20 ± 1.9%), followed by riverside (68.89 ± 2.75%) and urban areas (65 ± 2.8%) (Figure 6c,d). OM% of soil ranged from 1.7% to 2.74% which was relatively low but differed significantly (F = 7.79; p = 0.01) in all the habitats (Figure 6e).
Table 2. Descriptive statistics of the environmental variables affecting V. encelioides traits across different habitats in the region.
Figure 6. Box plots of the selected topographic, edaphic and soil physiochemical properties with statistically significant variations among the five habitat types (i.e., riverside (RS), cropland (CL), abandoned sites (AS), roadside (ROS), and urban land (UA); (a) Elevation (b) Latitude (c) Silt % (d) Sand % (e) Organic matter %.
Apart from topographic and edaphic factors, climatic variables also varied significantly between different habitats (Table 2). A higher DEW (4.1 ± 0.5), TMax (42–41.9) and TMean (19.2–18.7) and lower TMin (−3.8 to −4.49) were recorded at urban sites followed by riverside compared to other habitats. Factors such as SSW showed significant variation with comparatively high values at roadside (0.47 ± 0.01) followed by cropland. Several climatic variables differed significantly among habitats, including precipitation (F = 3.07; p = 0.02), summer drought severity index (F = 2.78; p = 0.03) and growing season precipitation (F = 3.2; p = 0.02), with high values recorded in abandoned land and low values in roadsides. In contrast, the relative humidity was higher in roadsides (47.5 ± 0.6) and lower in urban (43 ± 0.9) habitats.

3.3. Relationships Among Functional Traits and Habitats

The PCA biplot (Figure 7) and loading matrix (Table 3) show associations between plant functional traits and the studied habitats. The PCA explained a total of 44.7% variance, where the first axis explained 24.5% variation, and was clearly associated with seed properties (i.e., SN, SW, SL, SWd, and SDi) which were positively loaded on axis 1. The second axis explained 20.2% of the total variation and was linked to number of leaves, leaf area per plant, specific leaf area, inflorescence dry weight and vegetative dry biomass. On axis 3, average leaf length and width and leaf area index were positively loaded, whereas ALT was negatively loaded on both axis 1 and axis 2. Lower cumulative variance explained by the first two axes indicates the multidimensional trait variations distributed across additional PCA axes. Figure 7 suggests that seed mass traits (i.e., SW, SL, and SN) are closely correlated with each other and were associated with both cropland and riversides. The functional traits (average leaf length and width and leaf area index) were strongly correlated with each other and were found to be associated with urban sites. Number of flowers (NoF) was associated with abandoned land, where dendrometric parameters such as plant height (PHe) and stem diameter (SD) were inter-correlated and associated with road and abandoned land respectively.
Figure 7. Principle component analysis between the plant functional traits and different habitats of V. encelioides.
Table 3. PCA ordination results of plant functional traits.

3.4. Relationships Between Soil Variables and Habitats

The relationships between the habitats and environmental variables using RDA are shown in Table 4 and Figure 8. The results of the RDA explained 36.1% of the total variance. Some environmental parameters such as Asp (r = −0.37), Si% (r = −0.36) and pH (r = −0.33) were negatively correlated on axis 1. Conversely, Sa% (r = 0.31) and EC (r = 0.32) were positively loaded on axis 1 (Table 5). On axis 2, the dominant predictors were elevation (r = 0.30) and Lat (r = 0.34) which show positive associations, while DEW (r = −0.29), precipitation (r = −0.37), summer drought severity index (r = −0.40), and growing season temperature and precipitation (r = −0.39; r = −0.37 respectively) were negatively associated. On axis 3, Clay% and Lime%, were positively loaded while OM% and available nitrogen showed negative loading (Table 5).
Table 4. Results of the first three axes of redundancy analysis (RDA ordination).
Figure 8. Redundancy analysis (RDA) displaying the pattern of Verbesina encelioides functional traits along the environmental gradient.
Table 5. Correlation and biplot score of the first three axes of RDA.
The RDA biplot shows the pattern among the environmental variables and plant functional traits across different habitats (Figure 8). Key climatic factors (precipitation, growing season temperature and precipitation, summer drought severity index) negatively loaded on axis 2 show correlations with roadsides and riversides. EC and Lat loaded on the right side of the 2D ordination were predominantly influenced by urban areas and roadsides. However, factors like pH, Si% and Asp loaded on the left along axis 1 show associations with cropland. Si% on the left side of the ordination biplot also shows associations with riverside (Figure 8).

3.5. Hierarchical Cluster Analysis and Heatmap of Functional Traits and Environmental Variables Across Habitats

The hierarchical cluster analysis of the environmental variables and functional traits of V. encelioides is illustrated in Figure 9. The cluster analysis of the functional traits indicated that cropland and urban area were closely related, forming a cluster, while abandoned land and roadside habitats share similar seed characteristics and were grouped together. The riverside habitat forms a separate branch before joining the remaining clusters, indicating a distinct functional trait relative to the other habitats (Figure 9a). The heatmap showed that the populations growing in the riverside habitat showed a positive relationship with all functional traits except number of branches, while the roadside habitat showed a negative relationship except for number of flowers and average leaf thickness (Figure 9a). In Figure 9b, hierarchical clustering of environmental variables shows a different grouping pattern. Cropland and roadside are grouped together, while urban area and abandoned area are closely associated with each other (Figure 9b). In contrast, the riverside remains relatively distinct, showing different environmental conditions.
Figure 9. Hierarchical clustered heatmaps of (a) functional traits and (b) environmental variables across five habitat types. Note: Color intensity represents standardized (Z-score) values, while the dendrograms indicate the similarity among habitats based on hierarchical cluster analysis.
The heatmap of environmental variables represents that V. encelioides growing along the riverside shows a negative correlation with all soil physicochemical properties and topographic factors (except Sa% and longitude) while showing a positive correlation with all the climatic factors (except DEW and SSW). In contrast, abandoned land shows a positive correlation (except elevation, Asp, latitude, Si%, electric conductivity, TDS, bulk density and growing season temperature) with all environmental variables (Figure 9b).

3.6. Correlations Among Functional Traits of V. encelioides

The allometric relationships among the studied functional traits and biomass were examined using log-transformed linear regression models (Table S2); some traits showed significant correlation. The leaf variables like LAI × LAPP (R2 = 0.37), ALL× ALW (R2 = 0.44), and NoL × LAPP (R2 = 0.59) showed a moderate relationship compared to other traits (Figure 10a–c). However, a stronger relationship was observed among ALL × LAI and SLA × VDBM (R2 = 0.76) (Figure 10d,e). Other traits such as ALW showed a strong relationship with LAI (R2 = 0.86) (Figure 10f). The linear regression model showed significant variations in the reproductive traits of Verbesina encelioides, i.e., SN × SW (R2 = 0.65), SW × SL (R2 = 0.68) and SW × SWd (R2 = 0.71) (Figure 10g–i). Among the studied reproductive traits, SL showed a strong positive correlation (R2 = 0.93) with SWd (Figure 10j).
Figure 10. Linear regression model studying significant functional traits of V. encelioides. Note; (a) leaf area index × leaf area per plant; (b) alerage leaf length×average leaf width; (c) number of leaf × leaf area per plant; (d) average leaf length × leaf area index; (e) specific leaf area × vegetative dry biomass; (f) average leaf width × leaf area index; (g), seed number × seed weight; (h), seed weight × seed length; (i), seed weight × seed width; (j), aeed length × seed width.

4. Discussion

The present study demonstrates clear habitat-driven variation in functional traits of V. encelioides, highlighting its strong phenotypic plasticity across contrasting habitats. Plants growing along riverside habitats exhibited high values of vegetative growth (SD, LAI) and seed numbers. This may be related to greater climatic water availability and periodic water pulses in streams [59], which can promote plant growth and productivity, especially in invasive herbaceous species [60]. Riverside habitats may also provide greater water and nutrient availability, further supporting vigorous vegetative growth [61,62]. The changes in water availability, flooding and sediment conditions in riverside habitats create different microsites that influence the functional traits of invasive plants and support their growth and establishment [63]. In addition, floodwater may facilitate the long-distance dispersal of seeds, supporting the establishment and spread of V. encelioides along riversides and adjacent disturbed habitats. The observed habitat-related variation is consistent with the phenotypic plasticity of invasive plants, which allows them to adjust to variable environmental conditions across diverse habitats [64]. Previous studies have also reported that V. encelioides, as an invasive plant, may also form dense patches, particularly in disturbed habitats where suitable environmental conditions and local propagule availability facilitate establishment and promote further spread through competition for resources [65]. In addition, research from northern Pakistan indicates that altitude, soil nutrients, and soil texture are important environmental factors associated with communities invaded by invasive plant species [13]. In contrast, plants in abandoned areas showed increased numbers of branches and leaves and leaf area per plant but reduced plant height. These traits indicate the adaptive plastic responses of plants to environmental stress [66]. Such vegetative trait flexibility allows the plant to optimize resource use under contrasting conditions, thereby improving its survival and invasion success [67,68,69]. Abandoned lands, due to reduced management, may favor secondary succession and provide suitable conditions for the establishment, growth, seed production, and clustering of invasive plants [70]. Plants growing along roadsides displayed reduced ALL and ALW, but increased flower production. This trait reflects plant adaptation to higher light intensity, mechanical disturbance, or reduced soil moisture [71]. It is well established that plants in stressful environments typically produce smaller, tougher leaves to reduce water loss and resist mechanical damage [72]. In addition to these factors, roadside environments may expose plants to soil compaction and heavy metal pollutants which can affect root growth, resource availability and plant morphological traits [73]. The higher flower production of V. encelioides along the roadside may be linked to greater sunlight availability in these open habitats, as intense light conditions can enhance plant growth and reproductive output [74].
In cropland habitats, we found greater plant height and seed weight but a lower number of flowers. This pattern suggests a shift in resource allocation toward vegetative growth and the production of larger seeds under relatively favorable environmental conditions, consistent with life-history theory [75]. Larger seeds generally contain greater nutrient reserves, which can enhance seedling emergence, early establishment, and competitive ability in cultivated fields where crops compete intensely for light, water, and nutrients [76]. Our findings align with the widely recognized concept that habitat heterogeneity drives trait diversification in plant species [77]. Similarly, it was reported that increased plant height contributes to the invasive potential of V. encelioides in both cropping and non-cropping systems [30]. Taller plants are more effective at intercepting sunlight, and facilitate wider seed dispersal through wind beyond the surrounding vegetation, thereby enhancing colonization and establishment in new habitats [78].
Within each habitat, V. encelioides allocated more resources to vegetative and reproductive aboveground biomass than to belowground biomass. This pattern is common among many invasive species [79], as greater allocation to aboveground tissues can enhance light capture and competitive ability, potentially supporting colonization across different habitats [80]. Plants can also adjust biomass distribution according to resource availability to improve resource acquisition and use efficiency [81]. For instance, increased aboveground biomass may improve light capture and photosynthetic efficiency under low-light conditions [82,83]. Interestingly, relatively low biomass allocation was observed in cropland habitats, which may reflect stronger competition with cultivated or native plants that can influence resource availability and biomass production [84]. Despite differences in habitat conditions and soil nutrient availability (particularly nitrogen and phosphorus), no significant variation in biomass allocation was observed among the studied habitats. This relatively stable root-to-shoot allocation may indicate a degree of homeostatic resource allocation, whereby V. encelioides maintains a consistent investment between above- and belowground tissues while adjusting nutrient uptake and resource-use efficiency in response to local conditions. Such stability may enable the plant to balance investment between vegetative and reproductive growth, thereby supporting successful establishment and persistence across a wide range of environmental conditions [58].
We observed significant variations in the environmental conditions of five habitats under which the phenotypic responses of V. encelioides were evaluated. Cropland soils were characterized by higher organic matter and silt content, whereas the sandy soils of roadside, riverside and urban habitats suggested that variation in soil texture and nutrient availability enables the plant to exploit a wide range of environmental conditions and thereby enhance its invasion success [85]. The higher abundance of V. encelioides in sandy soils along roadsides and river margins may be associated with the good drainage and aeration of these soils, which can provide favorable conditions for the establishment and growth of some invasive plants, particularly in disturbed habitats [86,87]. High resource availability makes an area more vulnerable to invasion [88] than resource-poor areas [89,90]. However, some invasive plants can also thrive in low-nutrient habitats by employing resource conservation strategies, such as high resource-use efficiency [91]. Our findings are also consistent with reports of vigorous V. encelioides populations in the sandy soils of Oklahoma and Hawai’i, indicating that disturbed sandy habitats are particularly susceptible to invasion [35,92].
Significant differences were also observed in atmospheric variables, including Temp, DEW, SDSI, PPT and GSP, among the five habitats. In particular, habitats with higher temperatures and favorable moisture conditions supported greater growth and reproductive performance of V. encelioides. These findings are consistent with Ref. [93], in which the authors reported that increasing temperatures facilitate the establishment and spread of warm-adapted invasive species by creating less favorable conditions for many native plants. Similarly, higher precipitation can enhance soil moisture and nutrient availability, thereby promoting plant invasion [94]. The greater abundance of V. encelioides in lowland and midland habitats may therefore reflect the combined effect of suitable temperature, moisture availability, and soil conditions [30]. Our findings suggest that climatic, edaphic, and topographic factors interact to create favorable conditions for the establishment and spread of V. encelioides across contrasting habitats.
The morphological traits of V. encelioides revealed that individuals from cropland and river habitats had increased seed characteristics, which loaded positively on axis 1. This suggests an adaptation of the weed toward enhancing reproductive fitness [95]. These combinations of seed traits may allow V. encelioides to successfully establish itself in both sandy and silty soils. Seed architecture, therefore, plays an important role in adapting to different environmental conditions [96]. Moreover, seed morphology can also adjust in response to changes in wind velocity, particularly in high-elevation areas [97]. If a new habitat is unfavorable, the plant may adopt a strategy of reducing seed size while increasing seed number [98]. Conversely, higher values for leaf traits such as ALL, ALW and LAI loaded positively on axis 3 and were associated with urban areas. Human disturbance in urban areas can increase environmental heterogeneity, which further promotes increased resource availability or reduce competition from native species, creating opportunities for invasive plants to establish and spread [99,100]. Urban areas may also provide warmer microsites through the urban heat effect [101] while artificial irrigation can increase water availability during dry periods [102]. Together, these conditions may have the possibility to favor greater leaf development and LAI in V. encelioides. In addition, the larger leaves can enhance light interception and carbon assimilation and increase water loss, possibly representing a useful indicator of energy and water conservation [103,104].
Our RDA showed that sand percentage (Sa%) and electrical conductivity (EC) were positively associated with the measured traits of V. encelioides along axis 1, whereas silt percentage (Si%) and pH showed negative associations. These results suggest that V. encelioides is well adapted to sandy soils, where environmental conditions may favor its establishment and persistence. The association of soil texture, EC, and pH with V. encelioides traits suggests that the species can tolerate a range of soil conditions, while differences in pH may influence nutrient availability and water uptake. These findings suggest that the species has broad ecological tolerance, which may help it establish across diverse habitats. This finding is consistent with previous reports indicating that V. encelioides is a competent agricultural weed capable of invading disturbed habitats and reducing crop yields [105]. Additionally, Ref. [106] found dense populations of V. encelioides in Texas soils that were silty, slightly alkaline (pH 6.8–7.2), and low in organic matter (less than 1%). Interestingly, in Hawai’i, most populations have been found in coastal or disturbed areas near sea level, though the species has also been recorded at elevations as high as 2805 m [107]. This wide ecological range further highlights the remarkable adaptability of the species. Furthermore, silt-dominated soils provide favorable conditions for plant growth because of their high water-holding and nutrient retention capacity, with greater organic matter content that improves soil structure and fertility [108]. The key climatic factors (i.e., PPT, GSP, GST, and SDSI) were associated with roadside and riverside habitats and showed negative loading on axis 1. This reflects the high phenotypic plasticity and ecological adaptability of V. encelioides [109,110]. Similarly, Ref. [111] reported that climatic factors such as precipitation and temperature seasonality (i.e., annual precipitation, average temperature of the coldest quarter, and maximum temperature of the warmest month) were especially important in shaping the distribution of this species. In fact, the likelihood of V. encelioides occurrence increased with greater temperature seasonality and higher annual rainfall. Recently, V. encelioides was reported by [112] to be growing wild in local dry and arid habitats in Laurvara village in Banaskantha District, Gujarat. Similarly, Ref. [33] revealed that V. encelioides is a drought-tolerant annual plant and a cluster was recorded in Morocco, where it was found colonizing wastelands, roadsides, and croplands, which is in support of the observed ecological pattern in our study. Likewise, Ref. [113] observed that V. encelioides grows abundantly along roadsides, canal banks, and vacant areas, where it progressively displaces native plant species across the state of Punjab. These qualitative similarities in habitat preference further support the view that V. encelioides is a highly adaptable invasive species with the capacity to spread across a wide range of environmental conditions.
The hierarchical clustering of functional traits and environmental factors demonstrated that V. encelioides exhibits habitat-specific responses. The cropland and urban habitats are closely associated in terms of functional traits while abandoned land and roadside habitats are clustered together. These results suggest that similar resource availability, anthropogenic disturbances, and environmental conditions promote the functional trait plasticity and persistence of the invasive plant under frequent disturbance [114,115]. The observed clustering of environmental variables indicates that each habitat provides a different combination of soil and climatic conditions resulting from long-term anthropogenic influence [116] which influences the functional traits of V. encelioides.
The distinct separation of the riverside habitat, with its positive association with functional traits and climatic variables but negative correlation with most soil properties, suggests that moisture availability is a primary driver of functional trait variation and climatic factors exert a stronger influence than soil characteristics on functional trait expression in riverside environments [117]. The allometric analysis revealed that several leaf and seed functional traits of V. encelioides are closely related, reflecting combined patterns of plant growth and resource allocation. Similar functional coordination has been reported across a wide range of plant species, where leaf dimensions are closely associated with plant growth and productivity [118] reflecting the foliar economic spectrum through coordinated resource capture and investment in photosynthetic tissues. The strong positive relationships among seed functional traits suggest coordinated reproductive investment, consistent with the reproductive economic spectrum, where plants balance seed production and reproductive success under changing environmental conditions, potentially improving the fitness of invasive species [119,120]. Such functional trait integration shows the ecological adaptability and invasion success of V. encelioides across heterogeneous habitats. Compared with many native species, invasive plants generally exhibit greater phenotypic plasticity, enabling them to maintain growth and reproduction across a broader range of environmental conditions [121]. Our results are consistent with this pattern, suggesting that the phenotypic plasticity of V. encelioides contributes to its successful invasion in diverse habitats.
Study Limitations
The climatic variables used in this study were obtained from NASA gridded climate data for 1981–2021 because site-specific local climatic data were not available. Although these long-term data are useful for describing broad climatic patterns across the study area, they may not fully represent current-season microclimatic conditions, short-term rainfall events, or variations in temperature and moisture that can influence the growth and functional traits of annual plants. Future studies should use local station data that would help better explain the factors influencing the functional traits of Verbesina encelioides. The results suggest that disturbed habitats, especially roadsides and cultivated areas, should be prioritized for early detection and control of V. encelioides. Removing the plants before flowering and seed production, monitoring riverbanks after floods, and controlling plants in abandoned lands and croplands can reduce seed production, spread, and further establishment, supporting more effective habitat-specific management.

5. Conclusions

Our results revealed clear habitat-related variation in the functional traits of Verbesina encelioides. Riverside populations showed greater stem diameter, leaf area index, and seed production, while abandoned-land, roadside, and cropland populations showed distinct responses in branching, leaf traits, plant height, and seed traits. PCA and RDA further indicated that soil texture, EC, pH, and climatic variables were associated with variation in these functional traits. Sandy soils, particularly in roadside and riverside habitats, were associated with greater plant vigor, with higher flower production in roadside and stem diameter and LAI in riverside populations, suggesting that soil texture and related environmental conditions may favor growth and reproduction. The widespread occurrence of this species along riversides may also have been facilitated by the 2010 floods, which as the historical plausible facilitator created disturbed habitats and opportunities for seed dispersal and establishment. The contrasting trait responses among habitats suggest different ecological strategies, with greater branching and leaf production in abandoned habitats potentially reflecting responses to local disturbance and resource conditions, whereas greater vegetative growth along riversides may be associated with more favorable water availability. Roadside populations may promote the spread of V. encelioides by increasing seed movement and propagule pressure along transport routes, which is important to manage. The higher seed weight observed in cropland populations indicates greater reproductive investment, which may increase soil seed bank and competition with crops, potentially reducing its productivity and posing a risk to local food security. Environmental factors including soil texture, climate and topography further influenced plant traits and distribution patterns. Overall, disturbances associated with road construction, agricultural activities, vegetation removal, flooding, and land abandonment may create opportunities for invasion. These habitat-related responses link plant invasion to ecosystem disturbance and land-use change, making it important to monitor habitats to protect native vegetation and maintain ecosystem functions. Management should therefore be carried out with early physical removal of roadside plants before extensive flowering and seed maturation to reduce seed production and propagule pressure. In croplands, timely removal of plants before seed maturation and tillage practices can reduce soil seed-bank replenishment. Such habitat-specific management can help reduce invasion pressure while protecting biodiversity and supporting sustainable use of agricultural and natural land. Overall, early detection, reduction in seed production and propagule pressure, and restoration of disturbed habitats may improve long-term ecological resilience and contribute to the objectives of the Sustainable Development Goals. These measures can also support long-term restoration of ecosystems and biodiversity and more resilient land management in areas affected by invasive plants.

Supplementary Materials

The following supporting information can be downloaded at https://www.mdpi.com/article/10.3390/su18189305/s1, Table S1: Normality distribution using the Shapiro-Wilk test of functional traits of V. encelioides and Table S2. Allometric linear model of regression among functional traits of V. encelioides.

Author Contributions

A.K. and N.K. conceptualized the study design and conducted field and laboratory experiments; A.K. conducted formal analysis of the data and N.K. performed visualization and statistical data analysis; A.K. wrote the initial draft and N.K. reviewed and edited the manuscript; N.K. provided resources and supervision; M.A. performed review and language editing of the initial draft and gave suggestions; K.A. was responsible for funding acquisition and language editing. All authors have read and agreed to the published version of the manuscript.

Funding

This research was supported by the project “NatureLink: UK–Pakistan Regional Research Clusters for Biodiversity Monitoring and Climate Resilience” (Project No. 14818479), funded by the British Council.

Institutional Review Board Statement

Not applicable.

Data Availability Statement

All the raw data used in this research can be obtained from the corresponding author (nasrullah.uom@gmail.com) upon reasonable request.

Acknowledgments

The authors gratefully acknowledge Zainab for their valuable assistance and cooperation during the fieldwork. We also extend our sincere thanks to the Director of the Turnab Farm for their support and collaboration in the laboratory analyses. We also acknowledge the support of Rafi Ullah for their valuable suggestions and appreciation.

Conflicts of Interest

The authors declare no conflicts of interests.

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