Understanding the influence of precipitation and nitrogen depositions on the soil health and growth indices of invasive (Solidago canadensis L.) and native (Wedelia chinesis) association

Abstract

Climate change has impacted plant community sustainability and increased the risk of plant invasion. Variable precipitation patterns and nitrogen (N) deposition rates have influenced plant community structure and productivity in different ecosystems. These factors have important implications for ecosystem functioning and biodiversity in the face of climate change. The current study investigates the effects of precipitation levels and N deposition rates with various cropping system on soil and the growth indices of native WC (Wedelia chinesis (Osbeck) Merr.) and invasive SC (Solidago canadensis L.) plants. Two different pot experiments were conducted in a greenhouse. In the first experiment, three levels of precipitation (low precipitation (PL), normal precipitation (PN) and high precipitation (PH); while in N deposition experiment: three levels of N deposition (N₀ (no application), N₅ (5 g m^− 2 yr^− 1) and N₁₀ (10 g m^− 2 yr^− 1)) were evaluated with different cropping systems (bare pot as control (CK), invasive monocropping (SC), native monocropping (WC) and intercropping (SC ×WC)). Both experiments were executed in CRD-factorial design with five replications. During the experiment, soil chemical properties and growth index parameters were collected monthly. The precipitation experiment results showed that PL increased the soil NH₄⁺-N content in the native monocropping system, While, the maximum NO₃⁻-N content with application of PN as compared with control treatment (CK). Besides this, normal precipitation increased the total organic carbon (TOC) concentration with invasive monocropping by 59% and 70% during August and October month, respectively. Highest soil pH was recorded at combined application of normal precipitation and intercropping of native and invasive. The PN increased the growth index (chlorophyll content and plant height) and wet and dry biomass of shoots (53% and 45%) and roots (66% and 75%). Moreover, the maximum growth index and wet and dry biomass of shoots (49%) and roots (72% and 75%) were noticed in invasive monocropping cropping system. Furthermore, N depositions experiment results indicated that the addition of N (N₁₀) increased the concentration of NH₄⁺-N in intercropping of native and invasive plants during October and December. Similarly, the higher rate of N deposition (N₁₀) increased the soil NO₃⁻-N content in invasive monocropping during August and October. Whereas, appropriate application of N rate (N₅) in combination with invasive monocropping increased the TOC concentration by 44% in December. The addition of N₁₀ enhanced the growth index and wet and dry biomass of the roots (65% and 54%) and shoots (70% and 55%). Overall, PN, and increased of N deposition rate (N₁₀₎ favored the growth of invasive plants. This may increase the risk of plant invasion and have adverse impacts on native plants.

Supplementary Information

The online version contains supplementary material available at 10.1186/s12870-025-06814-1.

Introduction

It is generally accepted that climate change will substantially disrupt the natural and agricultural ecosystems [1]. The impact of global warming has been evident in the reduced temporal stability of plant community biomass [2], and is forecasted to potentially increase plant invasion [3]. Moreover, global climate change scenarios may advance biological invasion as warming trend generally favors biological invasion in many regions [4]. The risk of plant invasion is anticipated to rise in tandem with climate change [5]. Additionally, climate change is expected to change rainfall patterns [6], with projects indicating larger but less frequent rainfall events by the end of this century [7]. The influence of rainfall variability on plant community structure and productivity across different ecosystems remain uncertain [8]. Plants evolution in response to water availability has resulted in the development of specific traits [9]. Therefore, the variability in precipitation can significantly impact the function and structure of ecosystems [10]. This variability is characterized by dry and wet periods, where the increase in precipitation variability influences different plant species favoring during wet periods than dry periods. In times of increased variability, species with more profligate resource-use strategies tend to thrive by reducing water stress [11]. However, plant community responses to increased rainfall variability remain largely unclear [12].

The N is a key element required for plant growth and is one of the most important yield-limiting nutrients in crop production in all agro ecological regions worldwide. It is commonly taken up from the soil in one of two inorganic forms: NH₄⁺ and nitrate NO₃⁻ [13, 14]. Different N forms can affect the physiological and metabolic processes of plants, thus eventually influencing plant growth and crop yield [15]. The N deposition is one of the major climatic factors affecting plants [13, 16]. Over the past century, anthropogenic activities have greatly increased N deposition [17]. Invasive plants often invade ranges with a high level of resources, especially with high N levels [18]. The N deposition favors the growth and physiological advantages to non-native plants [19], worsening the plant invasion by decreasing the native shrub population [20]. This change in competition balance favors the fast-growing non-native plants.

Soil health refers to the continued capacity of soil to function as a vital living ecosystem that sustains plants, animals, and humans [21, 22]. This includes its ability to support plant growth, maintain environmental quality, and promote plant and animal health [23, 24]. Key indicators of soil health include nutrient availability (e.g., NH₄⁺-N and NO₃⁻-N content), organic matter content (e.g., TOC), soil pH, and overall soil structure [23, 25].

The impact of non-native plants on N cycling is multifaceted, influencing litter decomposition and altering the ratio of soil N forms [26, 27]. In contrast to native plants, non-native plants often exhibit more efficient nutrient utilization, potentially disrupting resource equilibrium and consequently affecting both plant biodiversity and overall ecosystem functions [28]. The specific preferences of certain chemical N forms may influence the distribution of plants and co-occurring species within a site [29]. Non-native plants that can monopolize soil N pools may suppress native plants directly or indirectly [30]. The invasion dynamics of non-native plants can be further catalyzed, especially if these non-native species are particularly susceptible to increased overall N availability. Additionally, their preference for specific N forms may hasten the proliferation of existing non-native plants [31, 32]. Wedelia chinensis (Creeping daisy) belong from family asteraceae is a perennial herb with sprawling growth, dense mats, simple opposite leaves, bright yellow flowers, fibrous root system and can grow up to 30 cm tall [33]. Wedelia chinensis is native to tropical and subtropical Asia, thriving in grasslands, forests and prefers moist and well drained soils [34]. Wedelia chinensis has ecological and medicinal importance by preventing soil erosion and use as anti-inflammatory, hepatoprotective and known for wound healing properties in traditional medicine [35]. In addition to this, it also has research significance, with insights into its response to environmental stressors like climate change [36].

Solidago canadensis L. (common goldenrod) belong from family Asterraceae is a perennial herb with upright growth, lance-shaped leaves, dense clusters, bright yellow flowers arranged in plume-like shape and rhizomatous root system [37]. S. candensis L. is native to North America, thriving in a variety of habitats, including open woodlands, meadow and prairies, and prefers full sun and well drained soils [37]. S. canadensis L. has ecological and medicinal importance by providing late-season nectar for pollinators such as butterflies and bees [37]. It traditionally uses as anti-inflammatory, diuretic and antimicrobial [38]. S. canadensis L. has research significance due to its potential invasive behavior, and its response to environmental changes [39].

The invasive plant, S. canadensis L., exerts significant adverse effects on the growth of native plants, particularly in hampering seed germination and seedling growth via the released allelochemicals [40]. Such inhibition of seed germination and seedling development plays a crucial role in successful invasion [41, 42]. S. canadensis L. disperses numerous small seeds easily carried by wind [43]. The process of plant invasion can be influenced by climate warming and atmospheric N deposition [44]. The extent of the synergistic effect on biomass production of S. canadensis L. varied with environmental conditions [45]. The invasive S. canadensis L. produced more biomass than the native plants under the same conditions. Interestingly, the proportion of biomass increased if the invasive S. canadensis L. is greater than that of the native plants under N addition treatment [46]. S. canadensis L. is considered an exceptionally successful invader worldwide largely due to its prolific vegetative and generative growth, making it a superior competitor over native species [47]. Invasive plant species usually grow faster, have a larger plant size, and accumulate more plant biomass than the species from the community in which they invade [48]. The greater aboveground biomass of S. canadensis L. is attributed to the larger shoot height and higher density of the plant, and these characteristics enable the S. canadensis L. plants to be more competitive [49].

In the presence of nutrient amendments and elevated temperatures, the invasive S. canadensis L. demonstrated superior performance compared to its native counterpart [44], and showcased a robust ability for asexual reproduction. The addition of N, coupled with interaction with increased temperature, led to increase in the biomass and morphological traits of S. canadensis L [50]. Alterations in biomass allocation patterns could potentially grant the invasive species a competitive advantage [51], especially considering the superior performance traits of invasive species over native ones [46]. Several studies have explored the impact of nutrients on S. canadensis L [49]. and its interaction with temperature [44, 52]. The present study aims to investigated the influence of precipitation levels and nitrogen deposition rates on the biomass and morphological traits of S. canadensis L. and Wedelia chinensis. In pursuit of this objective, the study focuses on addressing two key goals: (1) Assessing the impact of precipitation levels on soil health, growth indices, and competitive dynamics between invasive S. canadensis L. and native Wedelia chinensis within different four cropping systems. (2) Examining the effects of varying nitrogen deposition rates on soil nutrient dynamics and subsequent growth responses of invasive S. canadensis L. and native Wedelia chinensis, while considering the mechanisms of different competition and succession.

Materials and methods

Pot experiment, design and plant sampling

The pot experiments were conducted in the greenhouse at the Key Laboratory, School of Environmental Science and Safety Engineering, Jiangsu University, China (119°45′ E, 32°20′ N) from June to December in 2021. The natural conditions of the study location, where the invasive species S. canadensis L. is spreading, include a humid subtropical climate with average annual temperature 14.04 °C and annual precipitation of approximately 1000 mm. The growing season for S. canadensis L. typically spans from late spring to early autumn. The experiment was conducted using a Completely Randomized Design (CRD) with a factorial arrangement of treatments and five replicates (60 pots for each experiment) used in the current study. Two experiments were conducted separately for precipitation levels and N deposition rates with four cropping system. For the precipitation experiment, three levels of rainfall were used, normal rainfall (PN) (based on the monthly average rainfall of Zhenjiang City 109.65 mm), 20% increase in rainfall (PH) (132 mm), and 20% decrease in rainfall (PL) (88 mm) with four cropping system include invasive mono-cropping (SC), native mono-cropping (WC) and invasive + native intercropping system (WC × SC) and control treatment (CK) with neither invasive nor native plant in the pots. Both experimental plants were watered twice a week. The rainfall level for precipitation treatment was maintained using control sprinkle irrigation system.

Similarly, for the N deposition experiment, three levels of N were used: no N application as control (N₀), low N₅ rate (5 g m^− 2 yr^− 1), and high N₁₀ rate (10 g m^− 2 yr^− 1) in combination with control (CK); pot having no plant, invasive S. Canadensis L. mono-cropping (SC), native Wedelia chinesis mono-cropping (WC) and invasive × native intercropping system (WC × SC). Soluble nitrogen is the mixture nitrogen-containing inorganic salts as NH₄⁺-N: NO₃^–-N: CO(NH₂)₂-N as ratio (1:1:1).

Thus, 5 kg of well-prepared soil was weighed and placed in an open container as a nursery. Then, invasive and native seedlings of same sizes were transferred into pots with the planting design. After planting, all pots were cultivated in the greenhouse under natural conditions. Plants and soil samples were collected three times during the experiment period, in August, October, and December, for NO₃⁻-N, NH₄⁺-N, TOC, and pH. Similarly, plant height and weight were noted three times during the experimental period, and root length and weight measurements were taken when all the plants were harvested.

Material Preparation

Seeds of invasive S canadensis L. and native Wedelia chinensis were grown in two different tray pots in the greenhouse at the Key Laboratory, School of Environmental Science and Safety Engineering, Jiangsu University, China (119°45′ E, 32°20′ N). The experimental soil was collected from the surface soil (0–20 cm) of green space at the Jiangsu University campus (32°12′ N, 119°30′ E), Zhenjiang City, in March 2021. After removing the plant residues and stones, the air-dried soil was sieved through a 10-mesh (2 mm) and stored in the dark before use.

Plant growth and biomass parameters

The morphological parameters, including chlorophyll content, shoot length, wet and dry biomass were measured during August, October and December, 2021 year. A SPAD meter (SPAD-502, Minolta Camera Co., Osaka, Japan) was used to determine the SPAD values. Shoot length were measured using a ruler (cm), and an electronic digital caliper determined stem diameter (mm). The fresh weight of the plants, i.e., root and leaf fresh weight, was determined. Then, root and leaves were placed in an oven for 3 days at a constant temperature of 60 °C to determine the dry weights. Root fresh and dried weight was determined by carefully detaching the roots from the soil and washing them using distilled water to remove soil particles and other impurities. All biomass measures were taken (in g) using an analytical balance (Sartorius CP224S Balance, China).

Soil chemical parameters

To evaluate the soil NH₄⁺-N and NO₃⁻-N, samples from 0 to 10 cm depths were taken during August, October and December months 2021. Soil samples (wet soil) were passed through a 2-mm sieve. The samples were transferred to the polythene bag laboratory and placed in plastic bags to avoid any loss of moisture, and 5 g of each soil sample was extracted in 50 mL centrifuge tube, using 25 mL of 2 M KCl solution. NH₄⁺-N and NO₃⁻-N concentrations were determined using the spectrophotometer at wavelength 667 nm and 540 nm, respectively [53]. Soil pH was estimated in a 1: 5 soil: water (w/v) mixture using a glass electrode meter (PHS-3E, Shanghaileici, Shanghai, China). For total organic carbon (TOC) determination, 1 g of air-dried soil was taken in a 250 mL Erlenmeyer flask, added 10 mL potassium dichromate solution was mixed well (manually shaken), then added 20 mL concentrated sulfuric acid, shaken immediately; after oxidation reaction (Diluted with 100 mL of deionized water for about 10 min, shake well before filtration, and finally measured the absorbance value (610 nm) using a spectrophotometer.

Statistical analysis

Significant differences among treatments were tested using two-factor ANOVA followed by Duncans multiple comparisons tests, and the significant differences were accepted at P < 0.05. The ANOVA analysis was used to analyze the changes in soil chemical properties, plant growth parameters, and plant and root biomass among different treatments, and the mean values were compared. The effects of different precipitation levels and nitrogen deposition rates on soil nutrient and plant growth were analyzed with redundancy analysis (RDA) using CANOCO 5. All the statistical analyses were performed in IBM SPSS version 23.0 (SPSS Inc., Chicago, USA). The graph was drawn using Origin 2020.

Results

Soil chemical properties

Impact of precipitation levels and cropping system on soil NH₄⁺-N and NO₃⁻-N content

The impact of different precipitation levels and intercropping systems significantly affected soil NH₄⁺-N content in October, and December 2021 (P ≤ 0.05, Fig. 1). Specifically, the combined application of low precipitation and native mono-cropping significantly enhanced soil NH₄⁺-N by 32% in October and 31% in December compared to the control treatment (Fig. 1).

Fig. 1.

The precipitation levels on soil NO₃^–-N and NH₄⁺-N content in different cropping system. Lowercase letters indicate significant for the interaction of precipitation levels and cropping system variability (p < 0.05). Low precipitation (PL), normal precipitation (PN) and high precipitation (PH); control (CK); pot having no plant, invasive S. canadensis mono-cropping (SC), native Wedelia Chinesis mono-cropping (WC) and invasive + native intercropping system (WC × SC)

In an experiment investigating the effects of different precipitation levels and intercropping systems on soil NO₃⁻-N concentration, significant impacts were observed during the months of August, and October 2021 (P ≤ 0.05; Fig. 1). The interaction between normal precipitation and CK treatment significantly enhanced soil NO₃⁻-N concentration by 19%, and 21%, in August, and October, 2021 respectively (Fig. 1).

Impact of N deposition rates and cropping system on soil NH₄⁺-N concentration

The study investigated the effect of different rates of N deposition under different cropping system on soil NH₄⁺-N concentration (P ≤ 0.05, Fig. 2). The interaction of different N deposition rates and intercropping systems significantly affected soil NH₄⁺-N concentration (P ≤ 0.05; Fig. 2). The combined application of N₀ and intercropping (WC × SC) increased soil NH₄⁺-N by 40% in August. Moreover, the interaction of N₁₀ and intercropping of native and invasive species significantly enhanced soil NH₄⁺-N by 46% in October, while N₁₀ and CK increased NH₄⁺-N by 56% in December compared to other treatments.

Fig. 2.

The nitrogen deposition rates on soil NO₃^–-N and NH₄⁺-N content in different cropping system. Lowercase letters indicate significant for the interaction of nutrient levels and cropping system variability (p < 0.05). control (CK); pot having no plant, invasive S. canadensis mono-cropping (SC), native Wedelia chinesis mono-cropping (WC) and invasive + native intercropping system (WC × SC). Nº= no N application; N5 = normal N application; N10 = higher N application

The impact of different N deposition rates and cropping systems on soil NO₃⁻-N concentration. The study conducted measurements in August, and October of the 2021 year. The results revealed that both the N deposition rates and intercropping, as well as their interaction, had a significant effect on soil NO₃⁻-N levels, with a statistical significance level of P ≤ 0.05 (Fig. 2). Notably, the combined application of N₁₀ and native mono-cropping (N₁₀ × SC) exhibited a substantial increase (18%) in soil NO₃⁻-N concentration compared to other treatments in August. Additionally, the interaction between N₁₀ and CK significantly enhanced soil NO₃⁻-N levels by 21% and 20% in October and December, respectively (Fig. 2).

Impact of precipitation levels and cropping system on soil pH and TOC

The combined effects of different precipitation levels and cropping systems significantly influenced soil TOC concentration in August, October, and December 2021 (P ≤ 0.05, Table 1). The highest TOC values were recorded with normal precipitation and invasive mono-cropping. The precipitation level, cropping system and their interaction was significantly affected the soil TOC contents. The combined application of normal precipitation levels and the invasive mono-cropping system resulted in the highest recorded values of 16.3, 21.4, and 24.4 in August and October, respectively. These values surpassed those observed in other treatments. Similarly, soil pH increased notably under normal precipitation and intercropping of invasive and native weeds, with the highest values observed in August and October (Table 1). The combined effect of precipitation levels and cropping systems had a significant impact on enhancing soil pH throughout the three-month period. Notably, the highest soil pH values were recorded as 7.51 and 7.58 during the months of August, October, respectively, under the conditions of normal precipitation and intercropping of invasive and native weeds, surpassing the values obtained from other treatments. While application of low precipitation level and intercropping (SC × WC) demonstrated a significant improvement in soil pH by 7.58 compared to other treatments during December (Table 1).

Table 1.

Effect of precipitation and cropping system of invasive and native weeds on soil TOC content and soil pH

Treatments		TOC			Soil pH
Precipitation level (P)		Aug	Oct	Dec	Aug	Oct	Dec
	PL	8.29B	17.0 A	18.7B	7.29B	7.34B	7.45 A
	PN	10.1 A	17.7 A	20.3 A	7.37 A	7.43 A	7.37B
	PH	7.34 C	15.7B	19.1B	7.36 A	7.41 A	7.44 A
Cropping system (CS)
	CK	7.79B	16.4B	16.7 C	7.33	7.38	7.42AB
	SC	10.7 A	18.1 A	21.0 A	7.31	7.36	7.37B
	WC	7.65B	16.3B	19.9B	7.33	7.38	7.41AB
	SC × WC	8.16B	16.5B	20.0B	7.39	7.44	7.47 A
Interaction (P × CS)
PL	CK	6.97f	15.6ef	15.9	7.19e	7.24e	7.44bc
PL	SC	9.19b	18.0b	19.2	7.31 cd	7.36 cd	7.33 cd
PL	WC	8.14cde	16.9 cd	20.4	7.31 cd	7.36 cd	7.44bc
PL	SC × WC	8.85bc	17.5bc	19.5	7.35 cd	7.39 cd	7.58a
PN	CK	8.17bcde	17.0bcd	17.1	7.33 cd	7.39 cd	7.26d
PN	SC	16.3a	21.4a	24.5	7.27de	7.33de	7.38 cd
PN	WC	7.73cdef	16.4cde	19.4	7.38bc	7.43bc	7.38 cd
PN	SC × WC	8.22bcde	16.1def	20.3	7.51a	7.58a	7.45bc
PH	CK	8.22bcd	16.5cde	17.0	7.47ab	7.52ab	7.56ab
PH	SC	6.65f	15.0f	19.4	7.35 cd	7.40 cd	7.40c
PH	WC	7.08ef	15.6ef	20.0	7.30 cd	7.36 cd	7.41c
PH	SC × WC	7.41def	15.7def	20.1	7.31 cd	7.35 cd	7.39c
ANOVA	DF	P	P	P	P	P	P
P	2	32.0**	11.4**	16.7***	5.74*	6.80*	6.80*
CS	3	57.2***	19.8***	208**	2.78NS	2.50NS	3.10*
P × CS	6	56.5***	21.6***	45.7NS	7.04***	7.40***	5.20***

Lowercase letters indicate significant for the interaction of precipitation levels and cropping system variability (p < 0.05)0.2 Uppercase letters indicate variability between nutrient levels and cropping system (p < 0.05). low precipitation (PL), normal precipitation (PN) and high precipitation (PH); control (CK); pot having no plant, invasive S. canadensis mono-cropping (SC), native Wedelia chinesis mono-cropping (WC) and invasive + native intercropping system (WC × SC)

Impact of N deposition rates and cropping system on soil pH and TOC

The N experiment was conducted to examine the influence of different N deposition rates, cropping systems and their interaction on soil TOC concentration (Table 2). The interaction of N deposition rates and cropping systems significantly affected soil TOC content and pH throughout the study period (Table 2). The combination of N5 and invasive mono-cropping notably increased TOC and soil pH, particularly in August and December. The soil pH was significantly influenced by N rates and cropping systems, as well as their interactions, during the months of August, October, and December. The interaction between N₅ and invasive monocropping significantly enhanced soil pH, with increases of up to 7.56 and 7.03 observed in August and December, while interaction of N₅ and native enhanced the soil pH up to 7.03 in October respectively (Table 2).

Table 2.

Effect of nitrogen deposition application and cropping system of invasive and native weeds on soil TOC content and soil pH

Treatment		TOC			Soil pH
Nitrogen level (N)		Aug	Oct	Dec	Aug	Oct	Dec
	Nº	4.54 C	12.5 C	12.3 C	6.63 C	6.63B	6.63 A
	N5	8.37 A	16.8 A	18.3 A	7.32 A	6.90 A	6.65 A
	N10	5.26B	13.5B	13.2B	7.21B	6.83 A	6.55B
Cropping system (CS)
	CK	5.57B	14.1	14.5	7.10AB	6.79AB	6.57 C
	SC	6.77 A	14.8	15.3	7.03B	6.81 A	6.66 A
	WC	5.86B	14.0	14.2	6.94D	6.77B	6.61B
	SC × WC	6.03AB	14.1	14.4	7.14 A	6.78B	6.60BC
Interaction (N × CS)
Nº	CK	4.73	13.0	13.8d	6.96d	7.01ab	6.73bc
Nº	SC	5.49	13.5	12.9d	6.28 g	6.47f	6.55e
Nº	WC	3.80	11.8	11.3e	6.49f	6.26 g	6.72c
Nº	SC × WC	4.13	11.7	11.1e	6.80e	6.77 cd	6.53e
N5	CK	7.01	15.9	16.0c	7.08 cd	6.70de	6.28 g
N5	SC	9.55	17.4	19.9a	7.56a	7.04a	7.03a
N5	WC	8.21	16.8	18.4b	7.19bc	7.05a	6.48e
N5	SC × WC	8.71	17.2	18.8b	7.44a	6.83c	6.80b
N10	CK	4.98	13.4	13.7d	7.25b	6.67e	6.71c
N10	SC	5.26	13.4	13.1d	7.26b	6.99ab	6.40f
N10	WC	5.55	13.5	12.9d	7.15bc	6.93b	6.63d
N10	SC × WC	5.24	13.6	13.2d	7.18bc	6.73d	6.48e
ANOVA	DF	P	P	P	P	P	P
N	2	131***	136***	678***	150***	32.4***	5.96***
CS	3	3.63*	1.69NS	0.85NS	11.5***	3.10***	11.8***
N × CS	6	2.23NS	2.22NS	16.4***	31.8***	247***	219***

Note: 1 Lowercase letters indicate significant for the interaction of N rates and cropping system variability (p < 0.05)0.2 Uppercase letters indicate variability between nutrient levels and cropping system (p < 0.05). Nº= no N application; N5 = normal N application; N10 = higher N application; control (CK); pot having no plant, invasive S. canadensis mono-cropping (SC), native Wedelia chinesis mono-cropping (WC) and invasive + native intercropping system (WC × SC)

Impact on morphological parameters

Impact of precipitation levels and cropping system on chlorophyll content, plant height, shoot and root biomass

The chlorophyll content and plant height exhibited significant fluctuations influenced by precipitation levels, cropping systems, as well as their interactions throughout the months of August, October, and December (P ≤ 0.05, Table 3). The highest chlorophyll content and plant height were observed under normal precipitation and invasive mono-cropping. The interaction of precipitation levels and cropping system also significantly affected the chlorophyll contents. The highest chlorophyll content was recorded 39.6 when combined application of high precipitation and invasive monocropping was applied in August as compared to other treatments. Moreover, the maximum chlorophyll content was noticed 40.6 and 38.7 in October and December when combined application of normal precipitation and invasive mono-cropping was applied than other treated pots. Additionally, the highest pant height was observed the combined effect of normal precipitation and invasive weed on in August, compared to other treated soils. Additionally, in October and December, the highest plant height was recorded when normal precipitation and intercropped invasive weed were applied together, surpassing the growth observed in other treatments.

Table 3.

Effect of precipitation levels and cropping system of invasive and native weeds on plant height and biomass

Treatment		Chlorophyll			Plant Height			Wet Biomass		Dry Biomass
Precipitation level (P)		Aug	Oct	Dec	Aug	Oct	Dec	Shoot	Root	Shoot	Root
	PL	36.0B	36.4B	33.6B	7.18B	16.6B	23.8B	18.5B	21.7B	6.36 C	6.55 C
	PN	35.7 C	37.8 A	35.4 A	8.25 A	21.3 A	30.0 A	20.9 A	26.4 A	11.5 A	14.3 A
	PH	37.3 A	35.8 C	34.7 A	5.01 C	15.8 C	23.0 C	9.77 C	9.04B	10.5B	13.3B
Cropping system (CS)
	SC	38.9 A	37.0B	36.4 A	8.80 A	26.5 A	35.4B	18.7B	28.8 A	13.0 A	20.5 A
	WC	34.8D	37.8 A	34.0 C	5.68 C	11.5 C	16.6 C	28.6 A	23.2B	11.4B	11.7B
	I-SC	35.2 C	36.1 C	34.7B	7.41B	23.9B	36.3 A	8.55D	10.52 C	6.81 C	8.18 C
	I-WC	36.5D	35.7D	33.0D	5.36 C	9.63D	14.1D	9.61 C	8.06D	6.59 C	5.04D
Interaction (P × CS)
PL	SC	38.6b	35.5e	34.4c	9.89ab	25.9c	32.7d	26.3b	26.2c	9.09e	13.0d
PL	WC	34.4 g	38.8b	33.4d	5.82de	9.30i	15.3 h	24.9c	20.7d	6.14 h	3.85j
PL	I-SC	34.9 fg	35.8e	34.7c	7.64c	23.7d	35.4c	9.97 g	12.13f	5.18i	5.39i
PL	I-WC	36.2d	35.4e	32.0e	5.38ef	7.30j	11.7j	12.98e	10.83 g	5.03i	3.90j
PN	SC	38.4b	40.6a	38.5a	10.6a	27.6b	38.2b	17.0d	44.1a	17.2a	22.0b
PN	WC	33.7 h	36.8d	34.6e	6.90 cd	14.1f	21.5f	50.2a	42.3b	12.0d	20.2c
PN	I-SC	34.9f	36.8d	34.8e	9.10b	31.2a	42.1a	7.55ij	12.10f	7.90f	9.85f
PN	I-WC	35.7e	36.8d	33.7d	6.40cde	12.3 g	18.0 g	8.78 h	7.15hi	8.99e	5.09i
PH	SC	39.6a	35.0e	36.4b	5.90de	25.9c	35.2c	13.0e	16.1e	12.8c	26.4a
PH	WC	36.3d	37.7c	34.1c	4.33f	11.1 h	13.1i	10.86f	6.50ij	16.1b	11.2e
PH	I-SC	35.6e	35.6e	34.7c	5.50ef	16.9e	31.3e	8.13hi	7.32 h	7.34 g	9.31 g
PH	I-WC	37.6c	34.8f	33.4d	4.30f	9.30i	12.5ij	7.08j	6.19j	5.73 h	6.11 h
ANOVA	DF	P	P	P	P	P	P	P	P	P	P
P	2	416***	93.0***	68.9***	37.0***	1708***	768***	2232***	3100***	1376***	3011***
CS	3	781***	101***	130***	42.5***	4914***	3880***	5031***	5880***	811.6****	3851***
P × CS	6	14.4***	102***	20.6***	2.84*	215***	31.5***	2019***	2084***	152.0***	480.2***

Lowercase letters indicate significant for the interaction of precipitation levels and cropping system variability (p < 0.05). Uppercase letters indicate variability between nutrient levels and cropping system (p < 0.05). Precipitation (P), cropping system (CS), Precipitation × cropping system (P × CS), low precipitation (PL), normal precipitation (PN) and high precipitation (PH); control (CK); pot having no plant, invasive S. canadensis mono-cropping (SC), native Wedelia chinesis mono-cropping (WC), invasive intercropping (I-SC), native intercropping system (I-WC)

The influence of precipitation levels and cropping system on plant shoot and root biomass as shown in Table 3. Both the precipitation levels and cropping system and their interaction were significantly impact on the shoot and root biomass. Additionally, both fresh and dry shoot and root biomass were maximized under normal precipitation and invasive mono-cropping. Maximum fresh shoot and root biomass by 20.9 g and 26.4 g was found in normal precipitation level, while in cropping system, the maximum fresh shoot up to 28.6 by native monoculture cropping system, and invasive mono-cropping system enhanced the fresh root biomass increased by 28.8 g as compared with other treatments (Table 3). Moreover, the interaction of normal precipitation level and enhanced the fresh shoot biomass, while the maximum fresh root biomass by interactive effect of normal precipitation and invasive mono-cropping when compared with other treatments. According the precipitation experiment, the highest dry shoot and root biomass was noticed by application of normal precipitation. Additionally, in cropping system, the maximum dry shoot and root biomass was noticed by invasive mono-cropping system. However, the maximum dry biomass of shoot by the interactive application normal precipitation and invasive mono-cropping. Additionally, maximum dry root biomass by the interactive application high precipitation and invasive mono-cropping (Table 3).

Impact of N deposition rates and cropping system on chlorophyll content, plant height, shoot and root biomass

The interaction of N deposition rates and cropping systems and their interaction significantly influenced chlorophyll content, plant height, and biomass (P ≤ 0.05, Table 4). The interaction of N application and cropping system was also impact on chlorophyll content. The highest chlorophyll content was found when the combined application of N₅ and native mono-cropping during August, the maximum chlorophyll content was noticed with combination of N₁₀ and invasive plant of intercropping as compared with other treated soil in December. Moreover, the interaction of N rates and cropping system was also highly significant for plant height during October and December (Table 4). The maximum plant height was observed 21.1 and 31.1 cm when the combined application of N₅ rate and intercropped of invasive weed than other treatments in October and December (Table 4).

Table 4.

Effect of nitrogen deposition rates and cropping system of invasive and native weeds on plant height and biomass

Treatment		Chlorophyll			Plant Height			Wet Biomass		Dry Biomass
N deposition rate		Aug	Oct	Dec	Aug	Oct	Dec	Shoot	Root	Shoot	Root
	Nº	37.1 C	37.2	34.5 C	4.33 C	13.4B	16.7 C	10.8 C	8.77 C	6.29 C	6.63 C
	N5	39.7 A	37.8	36.0B	4.83B	15.5 A	21.5B	26.3B	21.6B	10.0B	13.4B
	N10	37.7B	40.4	39.3 A	5.45 A	15.8 A	22.3 A	35.6 A	25.4 A	14.0 A	14.5 A
Cropping system (CS)
	SC	39.5 A	40.4	36.7B	5.23B	17.7B	24.3B	36.9 A	25.3 A	12.4 A	13.3 A
	WC	37.4 C	37.9	36.7B	4.60 C	13.0 C	16.9 C	18.8 C	14.7D	11.0B	11.6 C
	I-SC	37.2 C	38.2	38.3 A	5.63 A	19.0 A	26.4 A	15.1D	17.0 C	7.3D	12.5B
	I-WC	38.4B	37.2	34.8 C	4.00D	9.90D	13.1D	26.07 C	17.5B	9.55 C	8.62D
Interaction (N × CS)
Nº	SC	36.8 g	38.6	35.4d	4.60	15.9c	20.0d	11.0i	6.00j	6.01 h	3.99i
Nº	WC	37.1f	36.9	33.5e	4.30	12.4e	15.2f	13.4 g	16.8f	9.17f	13.0e
Nº	I-SC	36.8 g	37.2	35.9d	4.80	15.6c	19.1e	10.0j	6.99i	5.21i	5.62 h
Nº	I-WC	37.4ef	35.9	31.3f	3.60	9.70f	12.5 h	8.84k	5.31j	4.75j	3.89i
N5	SC	42.6a	39.6	35.7d	5.30	18.6b	26.6c	47.5c	35.5a	12.6d	18.1a
N5	WC	39.0c	36.7	38.0bc	4.50	13.1d	17.0f	24.8d	18.1e	13.1c	13.8d
N5	I-SC	37.3f	38.0	37.5c	5.50	21.1a	31.1a	13.2 h	17.3f	7.16 g	15.0c
N5	I-WC	39.8b	36.7	34.7de	4.00	9.20f	11.4i	19.6f	15.4 g	7.01 g	6.74 g
N10	SC	39.7b	42.9	38.9b	5.80	18.5b	26.3c	52.1a	34.2b	18.6a	17.8a
N10	WC	36.0 h	40.1	38.5bc	5.00	13.5d	18.5e	18.2 g	9.29 h	10.8e	7.85f
N10	I-SC	37.6de	39.5	41.5a	6.60	20.3a	29.0b	22.1e	26.7d	9.59f	16.9b
N10	I-WC	37.9d	39.2	38.3bc	4.40	10.8e	15.4 g	49.8b	31.6c	16.9b	15.2c
ANOVA	DF	P	P	P	P	P	P	P	P	P	P
N	2	523**	352***	1110***	20.0***	352***	87.8***	3949***	3621***	3653***	9053***
CS	3	174***	57.0***	20.4***	88.9***	1115***	433***	3632***	999.4***	637.4***	621.1***
N × CS	6	108***	7.23***	3.41*	1.66NS	354***	5.03***	1787***	1414***	462.4***	1232***

Lowercase letters indicate significant for the interaction of precipitation levels and cropping system variability (p < 0.05). Uppercase letters indicate variability between nutrient levels and cropping system (p < 0.05). Nitrogen (N), cropping system (CS), nitrogen × cropping system (N × CS)Nº= no N application; N5 = normal N application; N10 = higher N application; control (CK); pot having no plant, invasive S. canadensis mono-cropping (SC), native Wedelia chinesis mono-cropping (WC), invasive intercropping (I-SC), native intercropping system (I-WC)

The impact of N deposition rates and cropping system on fresh and dry biomass of shoot and root as shown in Table 3. The maximum fresh and dry shoot and root biomass were observed at high dose of N application (N₁₀). In addition, cropping system, the maximum fresh and dry biomass of shoot and root were noticed by invasive monocropping system (P ≤ 0.05, Table 4). In addition, the maximum fresh and dry shoot biomass was noticed when combined application of N₁₀ and invasive mono-cropping was applied. Moreover, the highest fresh and dry root were observed with combined application of N₅ rate and invasive mono-cropping system (Table 4).

The PERMANOVA results showed considerable variances (p < 0.01) between the soil parameters and growth index (Fig. 3). The RDA results shown that the precipitation levels and cropping system recorded for 74% of the total variation (RDA1, 68%; RDA2, 6%) in soil parameters and growth index, wet-dry biomass (Fig. 3A). The RDA results revealed that the N rates and cropping system recorded for 79% of the total variation (RDA1, 76%; RDA2, 3%) in soil parameters and growth index, wet-dry biomass (Fig. 3A). The heatmap illustrates the correlations between the soil chemical properties, and growth index (plant height and chlorophyll content), wet-dry biomass (Fig. 4). The heatmap shows that changes in chemical soil properties influenced growth index biomass.

Fig. 3.

(A). Redundancy analysis soil chemical properties, growth index, and dry biomass at different precipitation levels and different cropping system. Redundancy analysis soil chemical properties, growth index, and dry biomass at different nutrients levels and different cropping system (B). A type III scaling (correlation) plot is shown, including all the constraining variables. The constraining variables explained 74% and 79% of the total variance of growth index and biomass. A taxonomic group arrow pointing in the same direction means a stronger positive correlation, and a longer arrow represents a stronger relationship. low precipitation (PL), normal precipitation (PN) and high precipitation (PH); control (CK); pot having no plant, invasive S. canadensis monocropping (SC), native Wedelia chinesis monocropping (WC) and invasive + native intercropping system (WC × SC). Nº= no N application; N5 = normal N application; N10 = higher N application

Fig. 4.

Show the correlation between the root and shoot biomass and plant growth index and soil nutrients for precipitation experiment (A); indicated the correlation between the root and shoot biomass and plant growth index and soil nutrients for nutrient experiment (B)

Discussion

Effects on soil nutrients

In present study soil NH₄⁺-N concentration differed slightly in all three precipitation levels during all three months. Notably, the highest NH₄⁺-N concentration in the soil was found during low precipitation during October within native mono cropping system, while in December, the highest NH₄⁺-N concentration were found in low precipitation in control treatment, followed closely by high precipitation in native mono cropping system (Fig. 2). These findings suggest that reduced precipitation can increase the concentration of NH₄⁺-N in the soil (Fig. 2), aligning with observations of [54], who reported increased NH₄⁺-N pools with reduced precipitation. This is attributed to reduced microbial biomass with decreased precipitation, leading to an increase in NH₄⁺-N [55]. Additionally, the decrease in precipitation may impede plant growth [56] and plant NH₄⁺-N uptake [57]. Microbial sensitivity to moisture changes further influences their activity, diminishing as soil moisture decreases, the consequently lowering rates of N cycling [55].

Examining the impact of precipitation levels on soil NO₃⁻-N concentration reveal an interesting pattern. The highest concentration was observed under normal precipitation, followed by high and then low precipitation. This underscores precipitation’s role in NO₃-N accumulation [58]. The increased NO₃⁻-N in normal precipitation can be attributed to optimal moisture levels facilitating the conversion of organic nitrogen to NO₃⁻-N [59]. Conversely, the slightly lower concentration of NO₃⁻-N in soil with high precipitation may result from leaching caused by excessive rainfall, potentially leading to NO₃⁻-N loss from the system [40]. The decreased concentration of NO₃⁻-N in soils with low precipitation may be due to limited water availability for the conversion of organic nitrogen to NO₃⁻-N [60]. It is important to note that these results are specific to the conditions and parameters of the study. Moreover [40], reported that precipitation intensification increased NO₃⁻-N concentration relative to the soil under control conditions. However, the nutrient level results indicate that the highest concentration of NO₃⁻-N was found in high N level (N₁₀) (Fig. 2). The cropping culture of precipitation experiment reveals that the maximum concentration of NO₃⁻-N was found in the bare pots (CK) soil compared to other treated pots (Fig. 2). In the cropping culture of nutrient experiment, the maximum concentration of NO₃⁻-N was found in invasive monoculture in August, while in October, the highest soil NO₃⁻-N concentration was in CK treated pot, respectively (Fig. 2). This may be due to plant uptake, with invasive plants exhibiting higher nutrient uptake [61]. In our experiment, the concentration of NO₃⁻-N is lower in high precipitation. This may be due to negatively charged ions, which can easily be leached from the soil profile with percolating water [62]. Another factor that might contribute to decreased soil NO₃⁻-N is the high rate of microbe denitrification under low precipitation. In denitrification, the microbes naturally convert NO₃⁻-N in the soil to gaseous N form. However, the pool of extractable NO₃⁻-N did not change with reduced precipitation [55].

The N deposition experiment slight differences in NH₄⁺-N concentration in soil across three levels, with high N levels exhibiting the highest NH₄⁺-N concentration (Fig. 1). Increased N fertilizer application is known to elevate soil NH₄⁺-N concentration, attributed to NH₄⁺ ions in the fertilizer that readily contribute to soil NH₄⁺-N levels [63]. The NH₄⁺ ions released are easily adsorbed by soil particles, influencing soil NH₄⁺-N levels [64]. The NH₄⁺-N undergoes transformations, including nitrification, influenced by factors like soil moisture and temperature [65]. Understanding the impact of higher N application on soil NH₄⁺-N is crucial for effective nutrient management and optimizing crop productivity [66]. The N₁₀ in this experiment likely contributed to increased shoot and root biomass, and also increased biomass for both root and shoot (Fig. 1). Maximum plant growth occurred at the N₁₀ rate followed by N₅ and N₀ rate (Table 4). The increased plant growth is attributed specifically to added NH₄⁺-N [67].

Soil nutrient availability plays a crucial role in the success of alien invasive plants, often exhibiting heightened nutrient use efficiency and adaptable utilization strategies compared to native species [68]. In nutrient-limited scenarios, invasive plants excel in carbon assimilation and nutrient acquisition, outcompeting natives. This advantage may stem from a resource conservative strategy, reducing nutrient requirements for sustained growth [69]. Invasive plants alter soil nutrient dynamics by releasing specific substrate compounds, modifying soil microbial communities, and creating an environment conducive to their growth [70]. Studies indicate invasive plants generate greater and superior quality leaf litter and root exudates, enhancing N and/or P availability, disrupting nutrient stoichiometry [71]. Changes in substrate availability contribute to shifts in soil microbial communities, accelerating nutrient cycling and release into the soil, establishing a self-reinforcing feedback mechanism [62].

Our results indicate that invasive species may utilize the increased availability of NH₄⁺-N and NO₃⁻-N in the soil, which is exacerbated by altered precipitation patterns. For instance, during periods of low precipitation, when native species are already challenged by reduced water availability, the invasive species’ superior nutrient uptake abilities [61], allow them to outcompete native species for the limited resources available. This competitive edge is further reinforced by the invasive species’ ability to thrive in high nutrient conditions, as evidenced by the highest NO₃⁻-N concentrations found in invasive monocultures (Fig. 2). Such conditions may not only favor the growth of invasive species but also enable them to modify the soil environment to their advantage, thereby suppressing native species growth [71].

Effects on soil TOC content and soil pH

The combined application of normal precipitation and invasive monocropping significantly increased soil TOC contents compared to the other treatments (Table 2). This interaction positively influences the accumulation of TOC in the soil, possibly due to moderate soil moisture enhancing faster litter fall and roots decomposition [72]. Moderate soil moisture may boost microbial activity and substrate transformation capacity [40], promoting litter and root decomposition, and increased plant-C transfer into mineral soil in nutrient treatment [62].

Invasive species, with robust growth and efficient nutrient uptake, likely contribute to higher organic matter inputs, increasing TOC content [73]. The combined treatment of normal precipitation and intercropping of native and invasive weeds resulted in the highest soil pH (Table 1) [74]. Soil pH is a critical soil property that affects nutrient availability and microbial activity [75, 76]. The interaction between normal precipitation and the presence of both native and invasive weeds in an intercropped system may have promoted enhanced nutrient cycling and altered rhizosphere processes, leading to a higher soil pH [77]. Additionally, the presence of invasive species could introduce unique root exudates or impact soil microbial communities, influencing pH [78].

We observed that sole application of N₅ rate and invasive mono-cropping significantly increased soil TOC contents (Table 1). This combination positively influenced organic carbon accumulation [79]. Nitrogen, essential for plant growth, at an appropriate rate like N₅, promotes productivity and contributes to increased organic matter inputs [40, 80]. Invasive mono-cropping may enhance nutrient uptake and organic matter decomposition, contributing to higher TOC content [81]. The sole application of N₅ and invasive mono-cropping also resulted in higher soil pH compared to other treatments. Soil pH, crucial for nutrient availability and microbial activity (Neina, 2019), may have been influenced by the interaction, affecting nutrient cycling, rhizosphere processes, and soil microbial communities (Table 1) [82]. It is noteworthy that N fertilizers, including N₅, can have alkalizing effects on soil pH.

Our findings revealed that the combined application of normal precipitation and invasive monocropping significantly increased soil total organic carbon (TOC) contents, pointing to the potential role of invasive species in enhancing organic matter inputs and influencing TOC accumulation. This interaction, possibly facilitated by moderate soil moisture, may have stimulated microbial activity and substrate transformation capacity, ultimately promoting litter and root decomposition and the subsequent transfer of plant-derived carbon into the mineral soil. Furthermore, the robust growth and efficient nutrient uptake exhibited by invasive species likely contributed to the observed increase in TOC content, underlining their capacity to influence soil organic matter dynamics (McLeod et al., 2021).

Effects on plant growth and biomass

In present study, the highest chlorophyll content and plant height were observed during normal precipitation level as compared with both low and normal precipitation conditions (Table 3). This underscores the vital role of optimal precipitation in fostering chlorophyll production and overall plant growth [83]. These conditions facilitate efficient water and nutrient absorption, leading to heightened chlorophyll synthesis and increased plant height [84]. Additionally, the direct effect of normal precipitation on root and shoot biomass was evident, as the plants exhibited greater biomass accumulation under these conditions. The adequate water supply during normal precipitation levels likely facilitates the development of a well-established root system, enabling efficient nutrient uptake and biomass production [85]. Moreover, reduced precipitation suppressed soil respiration and ecosystem photosynthesis, reflecting decreased aboveground biomass and productivity. This reduction limits nutrient availability due to water constraints on soil microbial processes [86]. Decreased precipitation not only suppresses plant biomass and physiological processes but can also cause mortality, as shown in a holm oak forest [87].

The choice of cropping system significantly influenced chlorophyll content and plant height, observed in different species. Both mono-cropping and intercropping of the invasive S. canadensis L. resulted in the highest chlorophyll content (Table 4). Additionally, the invasive mono-cropping system exhibited greater plant height compared to both native mono-cropping and intercropping of native and invasive weeds throughout the growing seasons [88]. Ecologists increasingly focus on invasion biology due to the increasing harm of plant invasions [48]. Success of invasive alien plants is attributed to broader environmental tolerance, higher phenotypic plasticity, and enhanced resource utilization capacity compared to native plants [89]. These traits provide adaptive advantages, contributing to their successful performance and competitiveness in new environments [90]. The invasive mono-cropping system offers favorable conditions conducive to greater plant height. The observed greater plant height in the invasive mono-cropping system may signal a potential threat to native species, as the invasive species may outcompete and suppress the growth of native plants, ultimately impacting the ecological balance and biodiversity within the ecosystem.

The N deposition experiment results demonstrated significant enhancements in plant growth (chlorophyll content and plant height and biomass) under high N deposition rate (N₁₀) followed by normal and control treatment (Table 3). High nutrient levels provide an abundance of essential nutrients, particularly N in the form of NO₃⁻N, promoting physiological functions crucial for plant development [91]. The N availability supports photosynthesis, protein synthesis, and overall metabolic activities [92]. With ample nutrients, plants allocate more energy to growth, leading to an increased plant growth index as a measure of overall vigor and productivity [40]. Moreover, high nutrient availability fosters greater biomass production in terms of both fresh and dry shoot biomass (Fan et al., 2020). The observed differences between high and normal nutrient levels emphasize the role of nutrient limitation in hindering plant growth and biomass accumulation [93].

In the cropping system, maximum plant growth index (chlorophyll content and plant height) observed in the intercropped invasive weed, while fresh and dry shoot and root biomass were highest in the invasive monocropped culture (Table 3). Competitive interactions in intercropping, where invasive weeds outcompete neighboring plants for resources, result in increased plant height and chlorophyll content [94]. The higher plant growth index in intercropped invasive weeds indicates their adaptability and thriving in mixed environments. Conversely, monocropped invasive cultures, free from resource limitations imposed by intercropping, exhibit even greater biomass accumulation due to traits enabling efficient nutrient uptake and aggressive root systems [95]. Observed differences are influenced by various factors, including specific invasive species, crop species, and management practices in the cropping system [96]. Consideration of invasive plants’ impacts on ecosystem dynamics and biodiversity is essential in evaluating their role in cropping systems.

Conclusion

This study highlights the considerable impacts of precipitation levels and N deposition rates in combination with different associations of native and invasive plant on soil chemical parameters and plant growth indices. The precipitation experiment results highlight that the interactive application of normal precipitation and invasive monocropping system had a great impact of on soil health (TOC, NO₃⁻-N and NH₄⁺-N) and improve the growth index (chlorophyll content, Plant height and wet-dry biomass). The study suggests that increased nutrient availability by normal precipitation contribute to the growth of invasive plants, which may pose risks to native plant populations. Furthermore, the N deposition experiment results indicated that combined application N₁₀ and invasive monocropping have positive effects on soil properties, plant growth and biomass. The findings also shed light on the growth patterns of invasive and native plant species under different nutrient conditions. These insights provide valuable knowledge for the management of ecosystems and conservation efforts, emphasizing the need for further research to deepen our understanding of these complex interactions and their implications for ecosystem resilience and biodiversity conservation.

Electronic supplementary material

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Author contributions

Ismail Khan: Investigation; Conceptualization; writing - original draft; Muhammad Tariq: Methodology, writing - original draft, review & editing, Conceptualization; Saleh Alfarraj and Sulaiman Ali Alharbi: Writing - review & editing; Investigation; Faisal Nadeem: Writing - review & editing, methodology; Muhammad Sadiq khan: Writing - review & editing, project administration, funding acquisition; Sezai Ercisli and Ayse Usanmaz Bozhuyuk: Writing - review & editing; Ping Zhuang: Conceptualization, project administration, funding acquisition. All authors have read and agreed to the published version of the manuscript.

Funding

This research was supported by Guangdong S&T Program (2022B1111230001) and Guangdong Provincial Key Laboratory of Applied Botany, South China Botanical Garden (2023B1212060046). The authors extend their appreciation to King Saud University, Riyadh, Saudi Arabia, for supporting this work through project (No. RSP2025R7).

Data availability

No datasets were generated or analysed during the current study.

Declarations

Ethics approval and consent to participate

Not applicable.

Consent for publication

Not applicable.

Competing interests

The authors declare no competing interests.