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Article

Priority Effects Favor Invasive Bidens frondosa over Its Native Congener Bidens biternata, While Late Arrival Incurs Higher Costs

Guangxi Key Laboratory of Plant Conservation and Restoration Ecology in Karst Terrain, Guangxi Institute of Botany, Guangxi Zhuang Autonomous Region and Chinese Academy of Sciences, Guilin 541006, China
*
Author to whom correspondence should be addressed.
Plants 2025, 14(16), 2515; https://doi.org/10.3390/plants14162515
Submission received: 26 June 2025 / Revised: 27 July 2025 / Accepted: 5 August 2025 / Published: 13 August 2025
(This article belongs to the Special Issue Plant Invasions and Their Interactions with the Environment)

Abstract

Priority effects—the phenomenon where early-arriving species influence the establishment, growth, and reproduction of later-arriving species during community assembly—play a critical role in determining the invasion success of exotic species. However, how priority effects are influenced by nitrogen (N) availability remains understudied. The invasive species Bidens frondosa has rapidly expanded its range in China over the past few years. Yet it remains unclear how priority effects in B. frondosa versus native species may mediate invasion success, as well as how nutrient levels may alter these effects. Addressing these questions is essential for understanding the mechanisms driving B. frondosa invasion and for developing effective management strategies. In a greenhouse experiment, we manipulated the planting order of B. frondosa and its native congener B. biternata, then measured the growth and competitiveness of B. frondosa and B. biternata in both control and N addition treatments. Planting order greatly impacted the growth and competitiveness of both B. frondosa and B. biternata. Early arrival had more positive effects on B. frondosa than B. biternata, while late arrival more strongly inhibited B. frondosa than B. biternata. For B. frondosa, priority effects lessened with nitrogen addition, but the opposite occurred for B. biternata. Thus, priority effects may favor B. frondosa invasion, while late arrival, particularly under nitrogen addition, may curb its spread. As such, sowing early-germinating native species represents a useful management strategy for controlling B. frondosa invasions.

1. Introduction

Invasive plant species frequently outcompete native plants, thereby reducing biodiversity. Understanding the mechanisms behind successful invasions is crucial for controlling invasive species and restoring affected communities [1,2]. Compared to their native counterparts, invasive plant species often exhibit higher resource capture and utilization capabilities, competitiveness, and reproductive rates [3,4,5]. Furthermore, invasive species often have earlier emergence dates, higher germination rates, and faster growth rates [6,7,8]. These characteristics may enable invasive plants to successfully establish themselves in new environments.
Priority effects represent a type of phenological niche separation, in which early-arriving species influence the establishment, growth, and reproduction of species that arrive later during community assembly [9,10,11]. Priority effects can influence plant species’ relationships and community composition [11,12,13]. Early-arriving species can reduce the resources (e.g., nutrients, water, and space) available for late-arriving species and thereby negatively impact their growth [14,15]. Alternatively, early-arriving species may modify the types of niches available for late-arriving species, such as via allelopathy; these modifications can have either positive or negative effects on later-arriving species [9,16]. Currently, most experimental studies examining priority effects (i.e., those manipulating the timing of species’ arrivals) have been conducted in temperate ecosystems, with far fewer studies of tropical and subtropical ecosystems [11]. The mechanisms underlying priority effects may depend on local environmental conditions and the relationships among species, as well as the ability of study species to adapt to environmental changes [17,18,19]. Plant phenology differs between tropical/subtropical climates and temperate climates, and the key factors influencing plant phenology also differ [20]. Whether the role of priority effects in tropical and subtropical ecosystems is different from that in temperate ecosystems remains unclear.
Priority effects are recognized as an important driver of invasion success for exotic species [21,22,23]. In most cases, priority effects provide greater benefits and impose fewer costs on invasive versus native species [11,24]. For example, Dickson et al. (2012) [25] found that priority effects can result in the formation of near-monocultures by invasive species, whereas early-arriving native species did not exhibit this tendency. Ferenc et al. (2021) [26] also found that arriving early significantly increases the biomass of alien species but not native species. However, this is not a universal finding, with other studies showing no difference in priority effects between invasive and native species [19,27]. In addition, Stuble and Souza (2016) [19] found that arriving late decreased the growth of native species more than invasive species. In contrast, a recent meta-analysis described significant reductions in the performance of invasive species that arrived late, but this was not seen for native species [23]. These discrepancies among studies might stem from differences among study species and systems [19]. To better understand how priority effects shape invasion success and resistance to invasion, both the potential advantages of arriving early and the disadvantages of arriving late must be explored across a wider array of invasive and native species, as well as study ecosystems [19].
The magnitude of priority effects can vary in response to environmental conditions [28,29]. For example, resource supplementation can strengthen priority effects, enabling early-arriving species to secure more resources [28]. In highly fertile soils, competitively inferior species may therefore achieve community dominance by arriving first [30]. Similarly, priority effects allow early-arriving species to gain additional resources and produce more biomass when soil nutrient levels are high but not when they are low [28]. Anthropogenic nitrogen (N) deposition increases the availability of soil N, a crucial nutrient for plant growth in natural ecosystems. Higher soil N levels due to human activities can therefore promote the growth of invasive species and enhance their competitive ability [5,31,32]. However, whether N addition also enhances the magnitude of priority effects for invasive species remains unknown.
Bidens frondosa, native to North America, has rapidly invaded natural ecosystems in China, where it is classified as a first-grade invasive plant species [33]. Aqueous extracts from B. frondosa have been shown to inhibit seed germination and seedling growth in Ageratum conyzoides and Plantago virginica [34]. Additionally, B. frondosa exhibits phenotypic plasticity in response to light, nitrogen, and water availability [35,36], produces a large number of seeds [4], and has a high germination rate [7]. In Guangxi Province of China, Bidens frondosa is sympatric with its native congener Bidens biternata, and both can be found in disturbed areas, such as rural roadsides and abandoned lands. Both species are annual herbs in the Asteraceae family and share similar flowering and seed-setting periods, typically from August to October. In subtropical areas, we have observed that the emergence of B. frondosa and B. biternata seedlings occurs from late March to early May. Species with similar functional traits are more likely to compete for resources [37,38]. However, it is unclear whether priority effects have contributed to the invasion success of B. frondosa, such as by positively affecting its growth and competitiveness.
In this study, we evaluated priority effects on the growth (i.e., total biomass, root/shoot (R/S) ratio, and reproductive ratio) and competitiveness (i.e., relative dominance index (RDI) and relative interaction index (RII)) of the invasive B. frondosa and its native congener B. biternata. We also examined how exogenous N addition impacted the strength of priority effects for both species. In our greenhouse experiment, we manipulated the timing of arrival via three planting order treatments: (1) T0—both species sown simultaneously; (2) T1B. frondosa sown three weeks before B. biternata (i.e., B. frondosa given priority); and (3) T2B. biternata sown three weeks before B. frondosa (i.e., B. biternata given priority). We simulated N deposition by adding N fertilizer to a subset of replicates.
We hypothesized that (1) priority planting will confer greater benefits on the invasive B. frondosa than on its native congener B. biternata; (2) arriving late will have less of a negative impact on the invasive species than the native congener; and (3) N addition will enhance the priority effects of B. frondosa to a greater extent than in the native congener.

2. Results

2.1. Growth Comparison of Invasive B. frondosa and Native B. biternata

Planting order (i.e., the timing of arrival), N addition, and their interaction significantly affected the total biomass of both B. frondosa and its native congener B. biternata (Table 1). For B. frondosa, as compared to T0 (both species sown simultaneously), the total biomass increased by 74.1% (with N addition) and 167.3% (without N addition) in T1 (B. frondosa given priority) (Figure 1a). The total biomass decreased by 72.7% in T2 (B. biternata given priority) under N addition, when compared to T0 (Figure 1a). B. frondosa grown without supplemental N produced 49.2% less biomass in T2 than in T0, though this difference was not statistically significant (Figure 1a). For B. biternata, as compared to T0, the total biomass increased by 73.4% (with N addition) and 17.7% (without N addition) in T2, but no significant differences were observed between T0 and T2 in the absence of supplemental N (Figure 1b). The total biomass decreased by 32.9% (with N addition) and 44.9% (without N addition) in T1, when compared to T0, though the differences were not statistically significant between T0 and T1 under N addition (Figure 1b).
Planting order and N addition both affected the R/S ratio of B. frondosa, but not that of B. biternata (Table 1). The R/S ratio of B. frondosa grown without supplemental N was greater in T2 (0.20 ± 0.01) than in T0 (0.14 ± 0.01) or T1 (0.15 ± 0.01) (Figure 1c). In addition, in T2, the R/S ratio of B. frondosa was higher in the absence of supplemental N (0.20 ± 0.01) than under N addition (0.15 ± 0.01) (Figure 1c). The R/S ratio of B. biternata did not differ among T0, T1, and T2 (Figure 1d).
Planting order affected the reproductive ratio of both B. frondosa and B. biternata, while N addition only affected the reproductive ratio of B. biternata (Table 1). The reproductive ratio of B. frondosa was lower in T2 (7.74 ± 0.72% with N and 6.8 ± 0.67% without N addition) than in T0 (14.63 ± 0.92% with N and 17.05 ± 2.19% without N addition) or T1 (16.45 ± 0.62% with N and 17.70 ± 2.25% without N addition) (Figure 1e). The reproductive ratio of B. biternata was greater in T2 (3.55 ± 0.32%) than in T1 (2.63 ± 0.17%) or T0 (2.42 ± 0.27%) under N addition (Figure 1f). In addition, in T2, the reproductive ratio of B. biternata was greater under N addition (3.55 ± 0.32%) than when unfertilized (2.60 ± 0.21%) (Figure 1f).

2.2. Competitive Performance of B. frondosa and B. biternata

Planting order affected both the RDI of B. frondosa and B. biternata (Table 1). As compared to T0, the RDI of B. frondosa increased by 61.3% (with N addition) and 123.2% (without N addition) in T1, but decreased by 74.5% (with N) and 42.8% (without N) in T2 (Figure 2a). For B. biternata, as compared to T0, the RDI increased by 52.4% (with N) and 17.4% (without N) in T2, but decreased by 43.1% (with N) and 50.1% (without N) in T1 (Figure 2b).
Planting order also affected the RII of both B. frondosa and B. biternata, while N addition only affected the RII of B. frondosa (Table 1). The RII of B. frondosa was greater in T1 (0.11 ± 0.4 with N and 0.00 ± 0.08 without N addition) than in T2 (−0.68 ± 0.05 with N and −0.66 ± 0.04 without N addition) or T0 (−0.17 ± 0.07 with N and −0.43 ± 0.05 without N addition) (Figure 2c). Under N addition, the RII of B. biternata was greater in T2 (0.29 ± 0.03) than in T1 (−0.21 ± 0.10) or T0 (0.03 ± 0.03). When no exogenous N was applied, the RII of B. biternata was also higher in T2 (0.11 ± 0.07) and T0 (0.05 ± 0.03) than in T1 (−0.24 ± 0.04) (Figure 2d).

3. Discussion

We examined priority effects on the growth and competitive performance of a subtropical invasive species, Bidens frondosa, and its native congener, Bidens biternata, in a greenhouse experiment including a nitrogen addition treatment. We found that early arrival conferred greater benefits to B. frondosa than to B. biternata, supporting our first study hypothesis. However, arriving late also incurred higher costs for B. frondosa than for B. biternata, contrary to our second study hypothesis.

3.1. Priority Effects on Growth in the Invasive B. frondosa and Its Native Congener

In our study, B. frondosa exhibited a greater percent increase in biomass when sown first as compared to B. biternata. This suggests stronger priority effects for B. frondosa than B. biternata, supporting our first study hypothesis. This finding is also consistent with several studies conducted in temperate regions, which showed that invasive plant species benefit more from priority effects than native species [24,25,26]. One possible explanation for this pattern is that invasive species generally have higher relative growth rates (RGRs) [39] and tend to grow rapidly in the absence of competition from other plants. In support of this explanation, in a previous study, we found that the RGR of B. frondosa was higher than that of B. biternata under favorable light and water conditions [40]. Having a higher RGR may enable B. frondosa to capture more light and ultimately grow larger than B. biternata.
Invasive plants often display rapid growth, strong competitive ability, and tolerance of resource-limited environments [41,42,43]; these characteristics may reduce the cost of arriving late and facilitate their successful establishment in native communities [19]. However, we found that arriving late decreased the biomass of B. frondosa to a greater extent than that of B. biternata, particularly under exogenous nitrogen addition. This suggests that B. frondosa incurred greater costs from arriving late than B. biternata. Priority effects are often mediated by niche pre-emption mechanisms, in which early-arriving species reduce the amount of resources (e.g., light, soil nutrients, and water) available to late-arriving species [9]. In an earlier study, we found that light levels significantly affect the growth of B. frondosa, which exhibits greater total biomass and a higher RGR under high versus low light conditions [40]. When planted first, B. biternata grew more vigorously, especially in the nitrogen addition treatment. This enhanced growth likely granted B. biternata a competitive advantage in capturing light resources, ultimately affecting light availability for late-arriving B. frondosa. We have also found that low light conditions reduce the RGR of B. frondosa to a greater extent than that of B. biternata [40]. Thus, the greater negative impact of arriving late for B. frondosa, as compared to B. biternata, may be due to reductions in the RGR under low light. This finding is inconsistent with those of previous studies in temperate regions, where arriving late imposed lower costs on alien species than natives [26]. This discrepancy among studies likely stems from differences in the study species and ecosystems [19].
Invasive plants may respond to changes in resource availability by altering biomass allocation patterns [44,45]. In the control (no N fertilizer), the R/S ratio of B. frondosa increased significantly when arriving late; this shift may enhance water and nutrient uptake under stressful conditions [46]. Bidens frondosa demonstrates high phenotypic plasticity in response to shifts in light, N, and water availability [36,40] as illustrated here. Additionally, arriving late reduced the reproductive ratio in B. frondosa. This may be related to a decrease in light availability, caused by the early arrival of B. biternata, which would have lowered the reproductive potential of B. frondosa. Ferenc et al. (2021) [26] also found that arriving late reduced the number of flowerheads in exotic species. Reductions in the reproductive ratio, along with a decrease in biomass, could negatively impact seed production, potentially lowering the invasion success of B. frondosa in the following season.

3.2. Priority Effects on Competitiveness in the Invasive B. frondosa and Its Native Congener

In community assembly, the timing of species’ arrivals significantly impacts competitive interactions between invasive and native species [47,48,49]. In this study, the RDI and RII of both B. frondosa and B. biternata decreased when planted late. This suggests that arriving first limited suppression by neighboring plants, thereby strengthening overall competitive ability. Our findings align with those of Stevens and Fehmi (2011) and Kardol et al. (2012) [28,47], who highlighted that species that arrive first are more likely to adapt to local environmental conditions and dominate the site [28,47]. Similarly, our results support the established notion that early-arriving species tend to outcompete those that arrive later [24,48,49]. Notably, when arriving early, B. frondosa showed a greater increase in RDI compared to B. biternata, suggesting that the priority effects for B. frondosa are stronger. Conversely, when planted second, the RDI and RII of both species decreased. This is consistent with previous studies that showed that the competitive ability of late-arriving species can be diminished by early-arriving ones [47,49]. Moreover, when planted second, the percentage decrease in RDI was more pronounced for B. frondosa versus B. biternata, and the RII of B. frondosa was lower than that of B. biternata. This implies that B. frondosa was less tolerant of competition from B. biternata when arriving late. Late arrival therefore incurred more significant competitive disadvantages for B. frondosa than B. biternata. This is consistent with previous findings from temperate regions, where early-arriving native species can competitively exclude invasive species [24,48].

3.3. The Magnitude of Priority Effects for Invasive B. frondosa and Its Native Congener Under N Addition

When exogenous N was applied, the magnitude of the beneficial effects of early arrival (i.e., enhanced biomass and RDI) decreased for B. frondosa but increased for B. biternata. This contradicts our third study hypothesis that N addition should amplify priority effects, particularly for the invasive B. frondosa. When B. frondosa is planted first, N addition may lessen the competitive inhibition of late-arriving B. biternata, increasing its growth and overall performance. This should weaken priority effects for B. frondosa under N addition. Several other studies have found that N addition may alleviate competition between invasive and native species [50,51]. When B. biternata is planted first, it may grow faster and larger when additional N is supplied, thereby more strongly shading late-arriving B. frondosa. This could decrease B. frondosa growth and thereby amplify priority effects in B. biternata. Similarly, early-arriving species gain access to additional resources in favorable environments, often leading to stronger priority effects in high- versus low-nutrient environments [28,49]. However, this is not a universal finding, and N addition does not always alter the strength of priority effects [52]. This may be because the strength of priority effects is mediated not only by impacts on resource levels, but also by the environmental sensitivity of early-arriving species and the overlap between competing species in terms of their resource needs [18]. Bidens biternata is a widespread weed in India and Japan [53,54], and it is capable of vigorous growth across diverse environments varying in light and water availability [40].
Although this study was conducted in a greenhouse, our results provide valuable insights into priority effects for the subtropical invasive species B. frondosa under different N regimes. Bidens frondosa and B. biternata are closely related phylogenetically, as well as being ecologically similar and sympatric in their distributions. Species with similar functional traits are more likely to compete for resources when sympatric [37,38]. Further research is needed to examine how other factors, such as water availability, moderate priority effects.

4. Materials and Methods

4.1. Study Site

This study was conducted at the Guangxi Institute of Botany (110°18′01.8″ E, 25°04′49.6″ N, 170 m a.s.l.), located in Yanshan, Guilin, Guangxi Province, China. This region features a subtropical monsoon climate, characterized by a mean annual temperature of 17.8 °C and an annual precipitation of 1742 mm. We conducted the planting order experiment in the Guangxi Institute greenhouse, under a natural photoperiod (temperature 25~35 °C and relative humidity 30~40%, recorded by a temperature and humidity meter, Beijing Honghai Yongchang Instrument Technology Development Centre, Beijing, China).

4.2. Plant Materials

In October 2016, we collected seeds from 20 Bidens biternata and 20 Bidens frondosa individuals growing on abandoned land within Guilin City, Guangxi, China. We stored the seeds in paper bags at room temperature. Both B. frondosa and B. biternata exhibit high seed germination rates, exceeding 90% and 75% at 20 °C, respectively [7,55].

4.3. Experimental Design

Our experiment was conducted in plastic pots with a diameter of 23 cm and a height of 18 cm. Each pot was filled with homogenized topsoil up to 5 cm from the top. The topsoil was obtained from an abandoned site in Guilin City. The baseline chemistry of the topsoil was as follows: the soil pH measured 7.14, organic matter 12.19 g kg−1, available N 117.63 mg kg−1, available P 29.45 mg kg−1, and available K 108.89 mg kg−1. To minimize the influence of the soil seedbank, the top 2 cm was removed before collecting topsoil to fill the experimental pots. Additionally, we regularly removed any non-target plant species (i.e., those not sown in the experiment) that emerged during the experimental period.
To establish priority treatments, we set a three-week sowing interval between B. frondosa and B. biternata, following the approach used by Ulrich and Perkins (2014) and Wilsey et al. (2015) [6,48]. For the simultaneous arrival treatment (T0), we sowed 10 seeds of each species into each pot on the same date. To establish priority for B. frondosa (T1), we sowed 10 B. frondosa seeds per pot on 28 March 2017, and allowed these to establish for three weeks. We then added 10 B. biternata seeds to each pot on 18 April 2017. The priority treatment for B. biternata (T2) followed the same procedure as for T1 but with the species reversed. Additionally, we sowed 20 seeds of each species separately in pots for use as monocultures. Plants were thinned to six plants per pot (three individuals per species in the mixtures and six individuals per species in the monocultures) after the emergence period concluded.
Nitrogen supplementation was initiated on 26 May 2017, when the seedlings of both species were well established. The average annual N wet deposition in China is 21.1 kg ha−1 year−1 [56]. As such, we tailored the N addition treatment so that experimental pots received 5 g/m2 N annually. NH4NO3 has been widely used to simulate atmospheric N deposition in many experiments [31]. Following He et al. (2012) [31], we applied N in the form of NH4NO3 dissolved in deionized water. A total of 0.645 g of NH4NO3 was added to each N addition pot on three dates (26 May, 3 June, and 12 June 2017). We also watered all pots daily throughout the experimental period. Each treatment included seven replicates.
In early August, we harvested all roots, shoots, leaves, and reproductive parts (i.e., inflorescences/peduncles, flowers, and fruits) from all experimental individuals; plant materials were dried at 70 °C until a constant weight and then weighed. We analyzed plant growth using the total biomass (the summed biomass of all roots, shoots, leaves, and reproductive parts), the root-to-shoot (R/S) ratio (root biomass/the summed biomass of all shoots, leaves, and reproductive parts), and the reproductive ratio (reproductive part biomass/total biomass).
We compared the competition performance of B. frondosa and B. biternata using the relative dominance index (RDI) [57] and the relative interaction index (RII) [58], which are based on the total biomass of each species. The RII was used as an estimate of the competitive response following Gruntman et al. (2013) [59]. We calculated the RDI and RII as follows:
RDIa = Bab/(Bab + Bba) × 100%
RDIb = Bba/(Bab + Bba) × 100%
RIIa = (BabBa)/(Bab + Ba)
RIIb = (BbaBb)/(Bba + Bb)
where a and b represent B. frondosa and B. biternata, respectively; Bab is the total biomass of B. frondosa when grown with B. biternata; Bba is the total biomass of B. biternata when grown with B. frondosa; Ba is the total biomass of B. frondosa in monoculture; and Bb is the total biomass of B. biternata in monoculture. The value of RDI ranges between 0% and 100%, and a high RDI indicates that the focal species is highly competitive [60]. The value of RII ranges from −1 to +1, with negative values indicating suppression by neighbors and higher (i.e., less negative) values indicating greater tolerance of competition [59].

4.4. Data Analysis

We analyzed the growth (total biomass, R/S ratio, and reproductive ratio) and competitive performance (RDI and RII) of B. frondosa and its native congener B. biternata separately. We used two-way ANOVAs to test for effects of planting order, N addition, and their interaction on growth and competitive performance in each species. We used one-way ANOVAs and least-significant difference (LSD) post hoc tests to evaluate the effects of sowing date in B. frondosa and B. biternata individually. For each species, an independent-sample t-test was used to compare samples with and without N addition. All tests were considered statistically significant at p < 0.05. We performed all statistical analyses using SPSS 18.0 (SPSS Inc., Chicago, IL, USA).

5. Conclusions

In our greenhouse experiment, arriving early enhanced the growth and competitive ability of B. frondosa to a greater extent than for B. biternata. Conversely, arriving late resulted in more pronounced reductions in growth and competitive ability for B. frondosa versus B. biternata. This suggests that B. frondosa experienced both stronger priority effects and also greater costs for arriving late. Furthermore, control and N addition treatments differed in terms of the shifts in biomass and RDI associated with planting order for both species. Therefore, the strength of priority effects is likely moderated by soil nitrogen availability. Our results illustrate how arrival timing influences B. frondosa invasion success, offering valuable insights for its control and the restoration of invaded plant communities. Arriving early may facilitate B. frondosa invasion, while, conversely, the early arrival of native species can suppress B. frondosa growth and competitiveness. Therefore, seeding early germinating native species represents a promising strategy to curb B. frondosa growth, thereby reducing invasion risk.

Author Contributions

Conceptualization, C.W. and S.T.; methodology, C.W. and S.T.; resources, C.W., X.L., Y.P. and L.Z.; writing—original draft preparation, C.W.; writing—review and editing, S.T.; project administration, S.T. All authors have read and agreed to the published version of the manuscript.

Funding

This research was funded by the National Natural Science Foundation of China (32260336), Natural Science Foundation of Guangxi Province (2025GXNSFAA069606), and Fundamental Research Funds for Guangxi Institute of Botany (23010).

Data Availability Statement

The original contributions presented in the study are included in the article; further inquiries can be directed to the corresponding author.

Conflicts of Interest

The authors declare no conflicts of interest.

Abbreviations

The following abbreviations are used in this manuscript:
RDIRelative dominance index
RIIRelative interaction index
RGRRelative growth rates

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Figure 1. Growth of Bidens frondosa (Left column) and Bidens biternata (Right column) under different planting order scenarios with supplemental N addition (N5) and without N addition (N0). (a) Total biomass of B. frondosa; (b) Total biomass of B. biternata; (c) Root/shoot ratio of B. frondosa; (d) Root/shoot ratio of B. biternata; (e) Reproductive ratio of B. frondosa; (f) Reproductive ratio of B. biternata. (T0: B. frondosa and B. biternata were sown at the same time; T1: B. frondosa was sown before B. biternate; and T2: B. biternata was sown before B. frondosa. Different lowercase letters (a, b) denote significant differences among T0, T1, and T2 without N addition, while different uppercase letters (A–C) indicate significant differences among T0, T1, and T2 with N addition; stars (*) indicate significant differences between N addition and control (no fertilizer) treatments for the same planting scenario, p < 0.05.)
Figure 1. Growth of Bidens frondosa (Left column) and Bidens biternata (Right column) under different planting order scenarios with supplemental N addition (N5) and without N addition (N0). (a) Total biomass of B. frondosa; (b) Total biomass of B. biternata; (c) Root/shoot ratio of B. frondosa; (d) Root/shoot ratio of B. biternata; (e) Reproductive ratio of B. frondosa; (f) Reproductive ratio of B. biternata. (T0: B. frondosa and B. biternata were sown at the same time; T1: B. frondosa was sown before B. biternate; and T2: B. biternata was sown before B. frondosa. Different lowercase letters (a, b) denote significant differences among T0, T1, and T2 without N addition, while different uppercase letters (A–C) indicate significant differences among T0, T1, and T2 with N addition; stars (*) indicate significant differences between N addition and control (no fertilizer) treatments for the same planting scenario, p < 0.05.)
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Figure 2. Relative dominance index (RDI) and relative interaction index (RII) for Bidens frondosa (Left column) and Bidens biternata (Right column) under different planting order scenarios with supplemental N addition (N5) and without N addition (N0). (a) RDI of B. frondosa; (b) RDI of B. biternata; (c) RII of B. frondosa; (d) RII of B. biternata. (T0: B. frondosa and B. biternata were sown at the same time; T1: B. frondosa was sown before B. biternate; and T2: B. biternata was sown before B. frondosa. Different lowercase letters (a–c) denote significant differences among T0, T1, and T2 without N addition, while different uppercase letters (A–C) indicate significant differences among T0, T1, and T2 with N addition; stars (*) indicate significant differences between the N addition and control treatments for the same planting scenario, p < 0.05.)
Figure 2. Relative dominance index (RDI) and relative interaction index (RII) for Bidens frondosa (Left column) and Bidens biternata (Right column) under different planting order scenarios with supplemental N addition (N5) and without N addition (N0). (a) RDI of B. frondosa; (b) RDI of B. biternata; (c) RII of B. frondosa; (d) RII of B. biternata. (T0: B. frondosa and B. biternata were sown at the same time; T1: B. frondosa was sown before B. biternate; and T2: B. biternata was sown before B. frondosa. Different lowercase letters (a–c) denote significant differences among T0, T1, and T2 without N addition, while different uppercase letters (A–C) indicate significant differences among T0, T1, and T2 with N addition; stars (*) indicate significant differences between the N addition and control treatments for the same planting scenario, p < 0.05.)
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Table 1. Two-way ANOVA of the effects of time of arrival (i.e., the planting order, T) and N addition (N) on growth and competitiveness in Bidens frondosa and Bidens biternata.
Table 1. Two-way ANOVA of the effects of time of arrival (i.e., the planting order, T) and N addition (N) on growth and competitiveness in Bidens frondosa and Bidens biternata.
SourcedfTotal Biomass
F-Value
R/S Ratio
F-Value
Reproductive Ratio
F-Value
RDI
F-Value
RII
F-Value
(a) B. frondosa
Time of arrival (T)247.04 ***6.90 **28.40 **100.50 ***82.74 ***
N addition (N)118.31 ***5.99 *0.620.886.69 *
T × N23.36 *2.020.723.122.87
(b) B. biternata
Time of arrival (T)232.47 ***2.195.23 **100.50 ***27.58 ***
N addition137.96 ***4.015.17 **0.881.63
T × N26.87 **0.022.933.121.63
Level of significance: * p < 0.05, ** p < 0.01, *** p < 0.001
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MDPI and ACS Style

Wei, C.; Tang, S.; Li, X.; Pan, Y.; Zhou, L. Priority Effects Favor Invasive Bidens frondosa over Its Native Congener Bidens biternata, While Late Arrival Incurs Higher Costs. Plants 2025, 14, 2515. https://doi.org/10.3390/plants14162515

AMA Style

Wei C, Tang S, Li X, Pan Y, Zhou L. Priority Effects Favor Invasive Bidens frondosa over Its Native Congener Bidens biternata, While Late Arrival Incurs Higher Costs. Plants. 2025; 14(16):2515. https://doi.org/10.3390/plants14162515

Chicago/Turabian Style

Wei, Chunqiang, Saichun Tang, Xiangqin Li, Yumei Pan, and Longwu Zhou. 2025. "Priority Effects Favor Invasive Bidens frondosa over Its Native Congener Bidens biternata, While Late Arrival Incurs Higher Costs" Plants 14, no. 16: 2515. https://doi.org/10.3390/plants14162515

APA Style

Wei, C., Tang, S., Li, X., Pan, Y., & Zhou, L. (2025). Priority Effects Favor Invasive Bidens frondosa over Its Native Congener Bidens biternata, While Late Arrival Incurs Higher Costs. Plants, 14(16), 2515. https://doi.org/10.3390/plants14162515

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