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Review

The Mechanisms of Sphagneticola trilobata Invasion as One of the Most Aggressive Invasive Plant Species

by
Hisashi Kato-Noguchi
* and
Midori Kato
Department of Applied Biological Science, Faculty of Agriculture, Kagawa University, Miki, Kagawa 761-1795, Japan
*
Author to whom correspondence should be addressed.
Diversity 2025, 17(10), 698; https://doi.org/10.3390/d17100698 (registering DOI)
Submission received: 26 August 2025 / Revised: 2 October 2025 / Accepted: 3 October 2025 / Published: 6 October 2025
(This article belongs to the Special Issue Ecology, Distribution, Impacts, and Management of Invasive Plants)
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Abstract

Sphagneticola trilobata (L.) Pruski has been introduced into many countries due to its ornamental and economic value. However, it has been listed in the world’s 100 worst alien invasive species due to its invasive nature. This species easily escapes cultivation and forms dense ground cover. It reproduces asexually through ramet formation from stem fragments. It also produces a large number of viable seeds that establish extensive seed banks. The movement of stem fragments and the dispersal of seeds, coupled with human activity, contribute to its short- and long-distance distribution. S. trilobata grows rapidly due to its high nutrient absorption and photosynthetic abilities. It exhibits high genetic and epigenetic variation. It can adapt to different habitats and tolerate various adverse environmental conditions, including cold and high temperatures, low and high light irradiation, low nutrient levels, waterlogging, drought, salinity, and global warming. S. trilobata has powerful defense systems against herbivory and pathogen infection. These systems activate the jasmonic acid signaling pathway, producing several defensive compounds. This species may also acquire more resources through allelopathy, which suppresses the germination and growth of neighboring plants. These life history traits and defensive abilities likely contribute to its invasive nature. This is the first review to focus on the mechanisms of its invasiveness in terms of growth and reproduction, as well as its ability to adapt to different environmental conditions and defend itself.

Graphical Abstract

1. Introduction

Sphagneticola trilobata (L.) Pruski, commonly known as wedelia, is a perennial herbaceous plant. It belongs to the Asteraceae family and was previously classified as Wedelia trilobata (L.) Hitchc. Its green, round, and solid stems easily root from their nodes. The leaves are simple and opposite. They are 4–9 cm long and 2–5 cm wide and are often three-lobed. The petioles are very short. The inflorescence is a capitulum. The capitulum is 2–3 cm in diameter and consists of 8–13 yellow ray florets and approximately one-hundred disc florets. The capitula are supported by 3–10 cm long pedicels that arise from the leaf axils. The fruits are brown achenes, or cypselae, measuring 3–5 mm in length. This species has a creeping and scrambling habit and forms dense, mat-like patches of ground cover [1,2,3,4,5,6,7,8] (Figure 1).
Sphagneticola trilobata is native to tropical America, from Brazil to Mexico, and the Caribbean [2,3,4]. It has been introduced into many countries as an ornamental plant. Due to its mat-forming habit and rooting stem nodes, it is useful for ground cover in landscaping and for controlling soil erosion [9,10,11,12,13]. However, it has easily escaped from its original planting areas and spread along roads and railways, to agricultural fields and other disturbed areas, and along forest edges. It has also been found in riparian and coastal vegetation. These areas include protected places in the tropical and subtropical regions of Australia, South Asia, East Asia, Southern Europe, Southern Africa, North America, and the Pacific Islands [2,3,7,13,14,15,16].
Dense ground cover patches of S. trilobata exclude other vegetation. The regeneration of native plant species is suppressed in areas covered by this species. Infestations of S. trilobata reduce the abundance and diversity of native plant species [2,3]. Due to its invasive nature, the International Union for Conservation of Nature (IUCN) has listed S. trilobata as one of the world’s 100 worst invasive alien species [17]. Additionally, a natural hybrid species between S. trilobata and its native congener Sphagneticola calendulacea (previously classified as Wedelia chinensis) was discovered in several locations in southern China in 2010 [18]. This hybrid species is highly competitive with S. calendulacea and can adapt to different environmental conditions [19,20,21,22,23]. Therefore, the hybrid species also threatens the conservation of the native species.
S. trilobata can be controlled through physical removal. However, this process must be repeated over an extended period of time because it can regrow from its underground parts. The species can also be controlled chemically by spraying 2,4-D, dicamba, glyphosate, and/or triclopyr [2,3,24]. Due to the survival of its underground parts and their re-sprouting nature, repeat chemical treatments are necessary. The ability of the species to sprout again from the remaining parts makes eradication difficult [2,3,24]. Currently, there is no specific control method available for this species. Long-term, continuous integrated management programs utilizing new technology may be necessary to eradicate S. trilobata.
Understanding the mechanisms of S. trilobata invasion is important for integrated weed management programs. Several other invasive plant species have been studied to understand their invasive mechanisms for this purpose. These traits include growth and reproductive capabilities, the ability to adapt to different environmental conditions, the ability to defend against herbivores and pathogens, and the ability to compete with neighboring plant species [25,26,27,28]. Information on these traits of S. trilobata has been accumulated over the past three decades. This review summarizes the invasive mechanisms of S. trilobata. We searched the literature using online databases, including Scopus, ScienceDirect, and Google Scholar. The following terms were searched in relation to S. trilobata or Wedelia trilobata: adaptation, allelopathy, botany, control, dispersal, diversity, flower, habitat, herbivory, impact, invasion, growth, mutualism, pathogen, reproduction, and stress tolerance.

2. Reproduction

2.1. Sexual Reproduction

S. trilobata produces capitula that bear achenes, or cypselae, year-round in tropical regions and under suitable conditions in other areas (Figure 1). These achenes contain seeds. However, the species is thought to spread primarily through asexual reproduction due to its limited fertile seed production [29]. Nevertheless, mature seeds have been reported in invasive populations of S. trilobata in Fiji and Japan [30,31,32]. On average, the Japanese populations produced 7.8 mature seeds per capitulum, resulting in a significant total seed production per plant. The extremely low self-pollination rate of 0.03% led us to conclude that cross-pollination was responsible. Their germination rate was over 80%. These seeds have spongy seed walls and hard seed covers. Thanks to their spongy seed walls, the seeds showed high buoyancy. After 90 days in seawater, the seeds remained viable due to their hard seed covers and buoyancy. As a result, the species has spread to the coastal areas of several islands in the Ryukyu Archipelago in Japan [30]. These characteristics of the seeds indicate their potential for long-distance dispersal in both seawater and freshwater. Populations of S. trilobata in Fiji produced a seed bank containing between 2100 and 3800 viable seeds per m2 [32]. Invasive populations of S. trilobata in China also produced mature seeds, suggesting sexual reproduction [33]. These findings suggest that S. trilobata produces a substantial number of viable seeds and establishes large seed banks, which contribute to population expansion through seed dispersal. Further research on the sexual reproduction of S. trilobata, particularly regarding mature seed production, is necessary in order to understand its role in population growth in its native and introduced ranges.

2.2. Asexual Reproduction

The stems of S. trilobata can grow to be over 2 m long. When these stems touch the soil, they easily develop nodal roots and sprouts, called ramets. These ramets then form genets, which are clone stands (Figure 1). Independent clonal plants form when these rooted nodes detach from the mother plants [2,3,34]. Stem fragments containing at least one node can also develop nodal roots and ramets, which produce new plants. Stem fragmentation of S. trilobata is often observed following disturbances, such as human activity and storms. Under field conditions, the average reproduction ratio from stem fragments was 90% [35]. Dumping garden waste also contributes to the dispersal of the fragments and to the expansion of these populations [2,3,36]. Therefore, asexual reproduction through fragmentation contributes to the growth of S. trilobata populations. S. trilobata is often found along riverbanks [8], which suggests that its stem fragments and/or seeds can potentially be transported by water flow. As described in Section 1, humans also facilitate the spread of this species overseas [9,10,11,12,13].

3. Growth Performance

S. trilobata populations have been observed to replace its native congener, such as S. calendulacea in southern China [13]. Several studies have compared the life history traits of the two species. S. trilobata has a relatively larger leaf area and biomass than S. calendulacea [37,38]. S. trilobata allocates more nitrogen to its photosynthetic apparatus [38]. Its photosynthetic capacity and photosynthesis-related gene expression are also higher. Additionally, S. trilobata contains higher levels of the plant hormones auxin and cytokinin, which contribute to the development of its large leaves [37].
Under different conditions, S. trilobata had greater shoot and root lengths, as well as greater above-ground and root biomass, compared to S. calendulacea. S. trilobata also had a greater nitrogen and phosphorus absorption ability than S. calendulacea [39,40,41,42]. S. trilobata contained higher levels of the plant hormone gibberellin. The concentration of gibberellin positively correlates with stem growth and the development of nodal roots from the stem nodes, resulting in active ramet formation in S. trilobata. Thus, gibberellin stimulates the development of the genets and the expansion of ground cover areas [43].
Endophytic bacteria of the Bacillus genus promote the growth of S. trilobata and the development of its genets but do not promote the growth of S. calendulacea [44]. Other endophytic bacteria, such as Sphingomonas, Pseudarthrobacter, and Novosphingobium, also promote the growth of S. trilobata [45]. S. trilobata alters the soil microbial population and stimulates microbial growth due to the high concentration of organic acids in its rhizosphere soil. This occurs through the root exudation of several organic acids, including propionic acid, fumaric acid, citric acid, acetic acid, and lactic acid. The decomposition of its residues also contributes to this process [42]. The decomposition rate of S. trilobata residues in the soil is approximately 40 days for 50% decomposition [46]. Altering the soil microbial population was found to promote the growth of S. trilobata [42].
These characteristics—higher photosynthetic capacity, higher nutrient absorption capacity, higher stem growth and genet formation, effective endophytic bacteria, and alteration of the soil microbial population—explain why S. trilobata grows faster than S. calendulacea. These characteristics may be associated with the invasive traits of this species. Additionally, the growth and expansion rate of the S. trilobata population were higher than those of other invasive Asteraceae plant species, such as Mikania micrantha, Bidens pilosa, Ageratum conyzoides, Aster subulatus, and Praxelis clematidea, in a man-made wetland in southern China [47].

4. Adaptation to Different Environments

4.1. Genetic and Epigenetic Variation

The chromosome number of S. trilobata is given as tetraploid, with x = 14 and 2n = 56 [48]. Genetic variation was high among S. trilobata populations from India when examined using inter-simple sequence repeat (ISSR) markers [49]. DNA methylation-sensitive amplified polymorphism (MSAP) and simple sequence repeat (SSR) analyses revealed significant genetic and epigenetic variation among S. trilobata populations from different regions of China. The correlation between epigenetics and geographic distance was stronger than the correlation between genetics and geographic distance [50]. S. trilobata populations on Hainan Island in China exhibited high phenotypic plasticity. The growth characteristics of this species, including stem length, number of internodes and branches, branching angles, and petiole angles, varied significantly among these populations. These characteristics were also altered by growth conditions, such as light intensity and plant density [51]. Therefore, S. trilobata exhibits high genetic and epigenetic variation. This variation could relate to greater species plasticity, enabling the species to adapt to different environmental conditions.

4.2. Habitat

S. trilobata thrives in riparian and coastal vegetation and along forest edges, roads, and railways. This plant also grows in agricultural fields and other disturbed areas in tropical and subtropical regions [2,3,7,13,14,15,16]. S. trilobata grows best in moist, well-drained, and fertile soil. However, it can also grow in clay, sand, or loam soils. The pH of these soils ranges from 5.5 to 7.5 [2]. S. trilobata is also tolerant of various abiotic stresses, including cold and high temperatures, low and high sunlight irradiation, low nutrition, waterlogging, drought, salinity, and global warming.
Mutualism with arbuscular mycorrhizal fungi (AMF) increases the levels of several amino acids, organic acids, flavonoids, and plant hormones in S. trilobata [52]. The accumulation of these compounds may enhance the ability of S. trilobata to respond to various stress conditions, enabling it to grow [52,53]. Consequently, S. trilobata grows in a wide range of habitats. Its mutualism with AMF suggests that it may be significantly impacted, which could contribute to its successful invasion.

4.3. Effects of Temperatures

S. trilobata thrives in an annual mean temperature range of 10 to 30 °C [2]. However, it can survive temperatures as low as 0 °C if it is acclimated at 15 °C. During this acclimation period, the levels of its total soluble sugars and anthocyanins increase, as does gene expression related to anthocyanin synthesis. Cold temperatures also generated reactive oxygen species (ROS) [54,55]. ROS can cause extensive damage to plant cells, including the photosynthetic apparatus, and may lead to death [56]. These anthocyanins illuminate ROS and protect the photosynthetic apparatus [57,58]. Increased levels of soluble sugars enhance cold tolerance in plants [59,60]. Therefore, the accumulation of anthocyanins and soluble sugars gives the plant cold tolerance. During cold stress, S. trilobata exhibits a higher photosynthetic capacity than its native congener S. calendulacea [54,55].
At a temperature of 35 °C, S. trilobata exhibited higher levels of photosynthetic pigments, chlorophyll fluorescence, and antioxidant systems that protect against ROS compared to S. calendulacea. Gene expression levels related to photosynthesis and antioxidant systems were also higher in S. trilobata [61,62]. Exposure to day/night temperatures of 40/35 °C and a heat wave of 40.1 °C results in higher growth rates and greater biomass production in S. trilobata than in S. calendulacea. This is due to the high photosynthetic activity of S. trilobata, which is achieved through its stable photosynthetic apparatus, and high water-use efficiency [61,62,63]. These findings suggest that S. trilobata possesses a more stable and efficient photosynthetic apparatus and antioxidant systems under high temperatures.
S. trilobata exhibits a greater photosynthetic capacity under cold- and high-temperature conditions. This is due to the thermostability of its photosynthetic apparatus and its ability to protect against ROS. It can also tolerate cold temperatures by accumulating soluble sugars.

4.4. Effects of Low Light Irradiation

S. trilobata prefers sunny habitats [2]. However, in both open and shaded environments (open: natural light; shaded: 25% of natural light), it exhibited higher growth rates in terms of biomass, stem length, and leaf number, as well as a greater photosynthetic capacity and water and nutrient use efficiency, compared to S. calendulacea [64]. S. trilobata maintains a relatively high photosynthetic ability and experiences less oxidative damage than S. calendulacea under shaded light conditions [65].
Similarly to what occurs under cold temperatures, S. trilobata accumulates anthocyanin under high light irradiation, turning its leaves from green to red [66]. Solar light energy is captured by chlorophyll pigments and transferred to photosystems I and II. These light-harvesting systems are located on the thylakoid membranes of the chloroplasts. Therefore, chloroplasts are the main cellular sites for ROS production. Anthocyanin illuminates ROS and protects cell structures, including photosynthetic apparatus [57,58]. Thus, S. trilobata can tolerate high light irradiation due to anthocyanin accumulation.

4.5. Effects of Nutrition

S. trilobata grows best in fertile soils [2]. However, under limited nitrogen conditions (0.91 mg/L with Hoagland nutrient solution), S. trilobata exhibited significantly higher growth, biomass, and chlorophyll content than S. calendulacea. Under these conditions, S. trilobata also exhibited significantly higher antioxidant defense systems, as indicated by the induction of catalase, peroxidase, and superoxide dismutase, compared to S. calendulacea [67]. These increased antioxidant defense systems may enable S. trilobata to grow at a higher rate under these conditions. S. trilobata also forms genets, by developing nodal roots and ramets from the stem nodes. Ramets that grow in high-nutrient conditions provide nutrients to the ramets that grow in low-nutrient conditions [68]. This integration of genets enables the plants to adapt to nutrient-heterogeneous environments.

4.6. Effects of Low Oxygen Stress

S. trilobata in riparian habitats often suffers from waterlogging conditions during high-water events, including flooding [69]. This results in low oxygen levels in the habitat. Plant species that are tolerant of low oxygen levels exhibit physiological and morphological adaptations, including the formation of aerenchyma [70,71,72,73]. Under these conditions, aerenchyma delivers oxygen, which contributes to the survival of these plant species [74,75]. Under waterlogging conditions, S. trilobata produced more aerenchyma than S. calendulacea [76]. S. trilobata exhibited higher growth, photosynthesis, and nutrient use efficiency rates than S. calendulacea [77,78]. These findings suggest that S. trilobata can tolerate waterlogging conditions. However, its physiological adaptations are unclear. Therefore, investigating these adaptations is necessary to better understand how the species tolerates these conditions.

4.7. Effects of Water Stress

S. trilobata grows in areas with annual precipitation ranging from 1000 to 2500 mm [2]. Drought conditions reduce its growth and photosynthetic activity. However, under these conditions, S. trilobata maintained relatively higher growth and photosynthetic activity than S. calendulacea [79]. Drought increases ROS production in the different cellular compartments of plants, causing extensive cellular damage [56]. S. trilobata increases the activity of superoxide dismutase, catalase, and peroxidase under drought conditions. These increases in enzyme activities are due to the rise in the concentration of the plant hormone abscisic acid (ABA) in the leaves [80,81,82]. These enzymes act as an antioxidant system by decreasing ROS in the cells and protecting them from damage [83,84]. An increased ABA level in the leaves also induces stomatal closure, which reduces transpiration in plants [80,81]. Therefore, S. trilobata may tolerate drought due to reduced ROS and transpiration through ABA induction.
High-salinity soils negatively affect plant vigor, photosynthesis, and growth [85,86]. High-salinity conditions also suppress the growth and biomass production of S. trilobata. However, S. trilobata showed relatively high growth and biomass production in salinity stress conditions of up to 100 mM of sodium chloride [87]. Soils with an electrical conductivity greater than 4 dS m−1 are classified as saline. This is equivalent to 40 mM of sodium chloride [88]. Therefore, S. trilobata exhibited relatively high tolerance to salinity stress. Salinity stress causes temporary water loss in leaf cells due to the osmotic effects of salts around the plant roots. The symptoms of water loss in leaf cells resemble those of drought stress. Salinity stress also induces high ROS production [85,86]. S. trilobata increases the activity of superoxide dismutase, catalase, and peroxidase under salinity stress conditions [89], which suggests that it enhances its antioxidant systems.

4.8. Global Warming

Global climate change includes an increased temperature and CO2 concentration in the atmosphere [90,91]. As discussed in Section 4.3, S. trilobata exhibits a higher rate of photosynthesis and growth at high temperatures. Its photosynthetic rate and total biomass increased by 67% and 70%, respectively, in an atmosphere with 700 μmol of CO2 compared to an atmosphere with 400 μmol of CO2. These increases are 2.3 to 2.7 times higher than those of S. calendulacea [92]. These results suggest an enhanced growth capacity in the context of rising CO2 levels. S. trilobata also exhibited greater increases in energy use efficiency and nutrient allocation to stem growth under elevated CO2 levels [93]. Therefore, S. trilobata thrives at high carbon dioxide levels and high temperatures, outcompeting the native species S. calendulacea. Consequently, S. trilobata may become more invasive and expand its distribution.

5. Defensive Response Against Herbivorous Insects and Pathogen

5.1. Herbivorous Insects

Herbivorous insects exert a significant selective pressure on the host plants, which suppresses their growth, survival, reproduction, and distribution [94,95,96,97,98]. Monophagous herbivores that are specific to respective invasive plant species are usually scarce in their introduced ranges due to a lack of coevolutionary history between the insects and these invasive plants [99,100]. However, invasive plants still face threats from oligophagous and generalist herbivorous insects in their introduced ranges. Many invasive plant species exhibit defense responses, and these defense responses likely contribute to their growth, survival, reproduction, and distribution [101,102,103].
In laboratory experiments, when S. trilobata and its native congener, S. calendulacea, were grown together, the larvae of generalist herbivorous insects, such as the cotton leafworm Spodoptera litura and the fall armyworm Spodoptera frugiperda, fed much more on S. calendulacea than on S. trilobata. Consequently, the growth reduction ratio due to herbivory was significantly higher for S. calendulacea than for S. trilobata [104]. These results suggest that S. trilobata has a higher antifeedant activity. Artificial defoliation treatments, which mimicked herbivory damage, suppressed the growth of S. calendulacea much more than that of S. trilobata [105]. This suggests that S. trilobata is more tolerant of herbivory damage. Additionally, when S. trilobata and other native plant species, such as Paederia foetida, Ipomoea nil, and Polygonum chinense, were grown in close proximity under field conditions, the larvae of S. litura and S. frugiperda fed much more on these native plant species than on S. trilobata [104]. This also suggests that S. trilobata has a higher antifeedant activity than these native plant species. The presence of S. trilobata causes increasing herbivory damage to nearby native plants. Therefore, S. trilobata has stronger antifeedant potential and greater tolerance to herbivory damage.
Aqueous extracts of S. trilobata leaves, stems, and/or flowers increased the mortality of the larvae of S. litura [106] and suppressed the oviposition activity of the fruit fly Bactrocera dorsalis [107]. Methanol, ethanol, and ethyl acetate extracts of S. trilobata leaves also increased the mortality of S. litura larvae, as well as those of the beet armyworm Spodoptera exigua and the diamondback moth Plutella xylostella [108,109]. Essential oil isolated from S. trilobata leaves and stems increased the mortality of the larvae of the red flour beetle Tribolium castaneum [110] and the adults of the maize weevil Sitophilus zeamais [111]. It also increased the mortality and anti-oviposition activity against adult cowpea weevils Callosobruchus maculatus [112]. These findings suggest that essential oils and extracts contain defensive compounds against herbivorous insects. These compounds may increase the mortality rate of herbivorous insects and reduce their oviposition activity. The major constituents of the essential oil were reported to be α-pinene (compound 1 in Figure 2; 18.2–34.9%), α-phellandrene (compound 2; 12.7–28.8%), limonene (compound 3; 4.5–17.7%), and germacrene D (compound 4; 54.4–15.8%) [110,111,112,113,114]. These monoterpenes (α-pinene, α-phellandrene, and limonene) and sesquiterpene (germacrene D) have been reported to act as defense compounds against herbivorous insects [115,116].
Following herbivory damage, the leaves of S. trilobata exhibited increased concentrations of flavonoids and jasmonic acid compared to those of native plant species, such as Paederia foetida, Ipomoea nil, and Polygonum chinens, under field conditions [104]. Certain flavonoids in other plant species have shown anti-herbivory activity [117,118]. Jasmonic acid is a signaling molecule that plays a role in defense against biotic stressors, including herbivory [119,120]. The caffeic acid ester derivative p-hydroxyphenyl caffeate (compound 8), which was isolated from the whole plant of S. trilobata, exhibited tyrosinase inhibitory activity with an IC50 value of 2 μM [121]. Tyrosinase is involved in tyrosine metabolism. This metabolism is responsible for immune responses and cuticle hardening in insects [122,123]. Therefore, inhibiting tyrosinase activity may hinder insect development and increase mortality.
The increase in insect mortality caused by extracts and essential oil indicates their insecticidal activity. Therefore, S. trilobata exhibits higher anti-herbivory activity, including antifeedant activity, anti-oviposition activity, and insecticidal activity. Jasmonic acid is known to increase the production of certain flavonoids, monoterpenes, sesquiterpenes, and diterpenes [124,125,126,127]. Therefore, S. trilobata may enhance its defensive ability against herbivory by increasing the production of flavonoids and terpenoids, such as α-pinene, α-phellandrene, limonene, and germacrene D, through elevated jasmonic acid levels. Additionally, p-hydroxyphenyl caffeate may also increase the defensive ability by disrupting tyrosine metabolism in insects. These defensive responses against herbivorous insects may contribute to the growth, survival, reproduction, and distribution of S. trilobata. However, the specific flavonoids involved in this defensive function have not yet been identified. Additionally, information on herbivory by mammals and other invertebrates is unavailable.

5.2. Pathogens

Pathogenic fungi, bacteria, and viruses also exert significant selective pressure on the host plants by suppressing their growth, survival, reproduction, and distribution [128,129,130]. The root exudates of S. trilobata inhibited the spore germination and growth of pathogenic fungi, such as Rhizoctonia solani, Fusarium oxysporum, and Phytophthora capsici [131]. R. solani infests a wide range of host plants and causes significant disease [132,133]. F. oxysporum and P. capsic cause Fusarium wilt syndrome and blight and fruit rot diseases, respectively, in a wide range of plant species [134]. The essential oil extracted from S. trilobata shoots was found to inhibit the growth of R. solani [135]. The major constituents of the essential oil of S. trilobata were described in Section 5.1. S. trilobata was also reported to produce more antimicrobial volatiles than that of S. calendulacea, including α-pinene (compound 1 in Figure 2) [136]. S. trilobata contains several ent-kaurane diterpenes, including 3α-angeloyloxy-16α,17-dihydroxy-ent-kauran-19-oic acid (compound 5), 3α-tigloyloxy-16α,17-dihydroxy-ent-kauran-19-oic acid (compound 6), and 3α-angeloyloxy-9β-hydroxy-ent-kaur-16-en-19-oic acid (compound 7). These compounds have been shown to exhibit antibacterial activity [137,138].
After being infected by R. solani, S. trilobata increased the expression of several genes, including an ethylene response factor within the jasmonic acid pathway. This resulted in increased jasmonic acid levels [139,140]. S. trilobata contains 3α-angeloyloxy-9β-hydroxy-ent-kaur-16-en-19-oic acid (compound 7). This compound activates the jasmonic acid signaling pathway, inducing defense functions against the gene expression of tomato spotted wilt virus (TSWV) and providing protection from TSWV infection [141]. Therefore, S. trilobata enhances the defense response to pathogen infection by activating the jasmonic acid signaling pathway. The mutualism of S. trilobata with the arbuscular mycorrhizal fungus (AMF) Glomus versiforme has also been reported to enhance defense against infection by R. solani [142]. Additionally, the evolutionary rate of pathogen resistance genes, such as TIR-NBS-LLR genes, is significantly higher in S. trilobata than in S. calendulacea [143]. Salicylic acid is another signaling molecule in plants. It is primarily responsible for defense against pathogen infection by producing defensive compounds, such as monoterpenes, sesquiterpene, and diterpenes [144,145,146]. However, the role of salicylic acid in the defense response of S. trilobata remains unclear. Clarifying the role of salicylic acid in the defense response against pathogen infection is essential.

6. Competitive Ability for Resource Acquisition

Competition among plants for resources, such as water, mineral nutrients, and sunlight, is often caused by neighboring plants. Allelopathy functions to suppress the germination and growth of neighboring plants by releasing allelochemicals into the surrounding environment, including the rhizosphere soil [147,148,149,150,151,152]. Consequently, plants that release allelochemicals have an advantage in accessing resources. Thus, allelopathy increases the completive ability of plants to acquire resources. Several invasive plant species have been reported to obtain more resources through allelopathy by suppressing the germination and growth of neighboring plant species [153,154,155,156]. These allelochemicals are produced and stored in various parts of the plants, including the leaves, stems, and roots, and are released when needed [157,158,159,160,161]. Therefore, certain plant parts contain allelochemicals, and these allelochemicals can be extractable.
Plant powder derived from the above-ground parts of S. trilobata was mixed with field soil, and the emergence of weeds was monitored. This resulted in the suppression of weed emergence. The degree of suppression depended on the concentration of the powder [162]. Aqueous extracts of S. trilobata leaves or litter inhibited the germination and growth of S. calendulacea [163], Amaranthus cruentus [164], and Eupatorium catarium [165]. These extracts also suppressed the germination and growth of several crop plants, including rapeseed (Brassica campestris), chickpea (Cicer arietinum), cowpea (Vigna unguiculata), mung bean (Vigna radiata), and common bean (Phaseolus vulgaris) [166,167,168]. Ethanol extracts obtained from the litter and rhizosphere soil of S. trilobata inhibited the germination and growth of lettuce (Lactuca sativa) [169]. These results suggest that S. trilobata contains certain allelochemicals in its above-ground parts and litter. Some of these allelochemicals are released into the rhizosphere soil. An essential oil derived from the above-ground parts of S. trilobata was found to inhibit the growth of weedy rice (Oryza, spp.) and lettuce [170]. The major constituents of the essential oil of S. trilobata, such as α-pinene, α-phellandrene, limonene, and germacrene D, have been reported to act as allelochemicals in many other plant species [171,172,173].
In addition, the chlorophyll content of Cyperus rotundus decreased with the increased application of aqueous leaf extracts of S. trilobata. The extract also increased the level of malondialdehyde and reduced the activity of peroxidase in C. rotundus [174]. The level of malondialdehyde in plant cells increases with oxidative stress. Oxidative stress causes cellular damage and death [56]. Conversely, peroxidase acts as an antioxidant system that reduces oxidative stress [83,84]. Therefore, leaf extracts from S. trilobata may induce oxidative stress and inhibit the antioxidant systems of C. rotundus. This disrupts the photosynthetic apparatus, including chlorophyll.
As described herein, S. trilobata exhibits allelopathic activity. The monoterpenes and sesquiterpenes found in its essential oil, including α-pinene, α-phellandrene, limonene, and germacrene D, may act as allelochemicals of S. trilobata. Extracts of its leaves, stems, litter, and rhizosphere soil may also contain allelochemicals. Some of these allelochemicals may be released into the rhizosphere soil. Through allelopathy, S. trilobata may obtain more resources by suppressing the germination and growth of neighboring plant species. This enables the species to grow and spread in the introduced ranges. However, the allelochemicals in the extracts and rhizosphere soils have not yet been identified. The allelopathic activity of the species in natural environments also remains unclear. In order to understand allelopathic function, it is necessary to identify these allelochemicals and determine their activity in the natural environment.

7. Conclusions

Based on the evidence we have compiled, S. trilobata has been introduced into many countries because of its ornamental and economic value. However, it easily escapes cultivation and forms dense, mat-like patches of ground cover. It spreads along roads and railways, into agricultural fields and other disturbed areas, and along forest margins. It has also been found in riparian and coastal vegetation. Its presence threatens the abundance and diversity of native plant species.
Table 1 summarizes the major mechanisms of S. trilobata invasiveness and includes the relevant references. This species develops ramets from the stem nodes, resulting in active genet formation. Stem fragments can also easily sprout ramets, which establish new plants and populations. The species is thought to spread primarily through asexual reproduction. However, the species has also been reported to produce a substantial number of mature seeds, which establish large seed banks. Seeds, stem fragments, and human activity contribute to the short- and long-distance distribution of the species. S. trilobata grows rapidly due to its high nutrient absorption ability, high photosynthetic ability, effective endophytic bacteria, and altered soil microbial population. Further investigation of its sexual reproduction, including the mature seed production, is necessary to understanding its role in population expansion.
S. trilobata exhibits high levels of genetic and epigenetic variation, which could contribute to its adaptation to different habitats and tolerance of various adverse environmental conditions, including cold and high temperatures, low and high light irradiation, low nutrient levels, waterlogging, drought, salinity, and global warming. The species tolerates these stress conditions through physiological and morphological adaptations, including the activation of antioxidant systems. Its mutualism with AMF also contributes to its growth and stress tolerance.
S. trilobata enhances the defense responses against herbivorous insects and pathogen infections by activating the jasmonic acid signaling pathway. Jasmonic acid induces the production of certain flavonoids and several terpenoids, including monoterpenes, sesquiterpenes, and diterpenes, as shown in Figure 2. These compounds may play a role in defense against pathogens and herbivory. The species also produces p-hydroxyphenyl caffeate. This compound inhibits insect development and increases insect mortality by inhibiting tyrosinase activity. S. trilobata exhibits allelopathic activity and produces several allelochemicals, including several monoterpenes and sesquiterpenes (Figure 2). Through allelopathy, the species may gain more resources by suppressing the germination and growth of neighboring plant species.
In summary, this species has a high reproductive capacity, grows rapidly, and exhibits a high level of genetic and epigenetic adaptability to different conditions. It also has defense mechanisms against herbivory and pathogens, as well as allelopathic properties. These characteristics may contribute to its survival and population growth in new habitats, establishing it as an invasive plant species. These documents may provide insight into effective weed management approaches. S. trilobata easily escapes cultivation and forms dense ground cover, suggesting that its invasive traits should be considered when handling it. Other invasive plant species listed among the world’s 100 worst alien invasive species [17], such as Mikania micrantha, Acacia mearnsii, Lantana camara, Arundo donax, Leucaena leucocephala, Reynoutria japonica, Imperata cylindrica, and Pueraria montana var. lobata, also exhibit a high reproductive capacity, rapid growth, high adaptability to different conditions, a high defensive ability against herbivory and pathogens, and allelopathic properties [26,28,102,153,154,155,175,176,177,178]. Therefore, these characteristics may be necessary to invasive plant species. Additionally, most of the available information on the invasiveness of S. trilobata is based on comparisons with its native relative, S. calendulacea. Further research is needed to understand the invasiveness of this species better. This could include comparisons with other native plant species.

Funding

This research received no external funding.

Informed Consent Statement

Not applicable.

Institutional Review Board Statement

Not applicable.

Data Availability Statement

Not applicable.

Conflicts of Interest

The authors declare no conflicts of interest.

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Figure 1. Sphagneticola trilobata. (A): Ground cover patch; (B): capitulum.
Figure 1. Sphagneticola trilobata. (A): Ground cover patch; (B): capitulum.
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Figure 2. Compounds involved in defense functions of Sphagneticola trilobata. Monoterpene: 1: α-Pinene; 2: α-Phellandrene; 3: Limonene. Sesquiterpene: 4: Germacrene D. Diterpene: 5: 3α-Angeloyloxy-16α,17-dihydroxy-ent-kauran-19-oic acid; 6: α-Tigloyloxy-16α,17-dihydroxy-ent-kauran-19-oic acid; 7: 3α-Angeloyloxy-9β-hydroxy-ent-kaur-16-en-19-oic acid. Caffeic acid derivative: 8: p-Hydroxyphenyl caffeate.
Figure 2. Compounds involved in defense functions of Sphagneticola trilobata. Monoterpene: 1: α-Pinene; 2: α-Phellandrene; 3: Limonene. Sesquiterpene: 4: Germacrene D. Diterpene: 5: 3α-Angeloyloxy-16α,17-dihydroxy-ent-kauran-19-oic acid; 6: α-Tigloyloxy-16α,17-dihydroxy-ent-kauran-19-oic acid; 7: 3α-Angeloyloxy-9β-hydroxy-ent-kaur-16-en-19-oic acid. Caffeic acid derivative: 8: p-Hydroxyphenyl caffeate.
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Table 1. The mechanisms of Sphagneticola trilobata that likely enhance its invasiveness.
Table 1. The mechanisms of Sphagneticola trilobata that likely enhance its invasiveness.
CharacteristicReferences
High reproduction
⮚
Ramet formation from stems and its fragments.
[2,3,8,34,35,36]
⮚
Substantial mature seed production and large seed banks.
[30,32,33]
Rapid growth and active genet formation
⮚
Higher photosynthetic capacity.
[37,38]
⮚
Higher nutrient absorption capacity.
[39,40,41,42]
⮚
Effective endophytic bacteria and alteration of soil microbial population.
[42,44,45]
High adaptative ability to different conditions
⮚
High genetic and epigenetic variation.
[49,50,51]
⮚
Mutualism with arbuscular mycorrhizal fungi.
[52,53]
⮚
Tolerance to cold and high temperature.
[54,55,61,62,63]
⮚
Tolerance to low and high light irradiation.
[64,65,66]
⮚
Tolerance to low nutrition.
[67,68]
⮚
Tolerance to waterlogging.
[76,77,78]
⮚
Tolerance to drought and salinity.
[79,80,81,82,87,89]
⮚
Tolerance to global warming.
[92,93]
High defense ability against herbivory, pathogens, and competitive plant species
⮚
Anti-herbivory activity.
[104,105,106,107,108,109,110,111,112]
⮚
Anti-pathogen activity.
[131,132,133,134,135]
⮚
Induction of the jasmonic acid signaling pathway.
[104,139,140,141]
⮚
Production of defensive compounds.
[110,111,112,113,114,115,116,121,136,137,138]
⮚
Allelopathy and production of allelochemicals.
[162,163,164,165,166,167,168,169,170,174]
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Kato-Noguchi, H.; Kato, M. The Mechanisms of Sphagneticola trilobata Invasion as One of the Most Aggressive Invasive Plant Species. Diversity 2025, 17, 698. https://doi.org/10.3390/d17100698

AMA Style

Kato-Noguchi H, Kato M. The Mechanisms of Sphagneticola trilobata Invasion as One of the Most Aggressive Invasive Plant Species. Diversity. 2025; 17(10):698. https://doi.org/10.3390/d17100698

Chicago/Turabian Style

Kato-Noguchi, Hisashi, and Midori Kato. 2025. "The Mechanisms of Sphagneticola trilobata Invasion as One of the Most Aggressive Invasive Plant Species" Diversity 17, no. 10: 698. https://doi.org/10.3390/d17100698

APA Style

Kato-Noguchi, H., & Kato, M. (2025). The Mechanisms of Sphagneticola trilobata Invasion as One of the Most Aggressive Invasive Plant Species. Diversity, 17(10), 698. https://doi.org/10.3390/d17100698

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