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Article

The Addition of an Invasive Plant Alters the Home-Field Advantage of Native Leaf Litter Decomposition

by
Shaojun Chen
1,*,
Xiaohua Xie
1,
Jie Wen
2,
Hao Zhai
1,
Huiqi Wang
1,
Yuhang Jiang
1 and
Zhanxu Gou
1
1
School of Green Intelligence Environment, Yangtze Normal University, Chongqing 408100, China
2
State Key Laboratory of Simulation and Regulation of Water Cycle in River Basin, China Institute of Water Resources and Hydropower Research, Beijing 100038, China
*
Author to whom correspondence should be addressed.
Forests 2024, 15(10), 1708; https://doi.org/10.3390/f15101708
Submission received: 27 August 2024 / Revised: 20 September 2024 / Accepted: 25 September 2024 / Published: 27 September 2024
(This article belongs to the Special Issue Adaptation of Forest Lichen and Moss Systems to Invasive Species and Disturbances)
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Abstract

:
Forest litter can decompose faster at home sites than at guest sites (home-field advantage, HFA), yet few studies have focused on the response of the HFA of native plant decomposition to the presence of invasive plants. We loaded the dry leaves of native Neosinocalamus affinis (decomposition resistant) and Ficus virens (more easily decomposable) leaves into litterbags with and without invasive Alternanthera philoxeroides, and incubated these litterbags at N. affinis and F. virens sites at the edge of the forest. The results showed that positive HFA effects with litter mass loss were at least 1.32% faster at home sites than at guest sites. The addition of A. philoxeroides reduced the mean HFA of N. affinis litter and increased that of F. virens litter. The HFA index without A. philoxeroides was significantly higher than that with A. philoxeroides. Soil faunal abundance colonized at home sites was always higher than that colonized at guest sites. Compared with the F. virens site, the abundance of Collembola, Arachnida, Formicidae and Lepismatidae at the N. affinis site was significantly higher compared to the F. virens site, while the abundance of Isopoda, Oligochaeta, Nematoda and Dermaptera was significantly lower. Our results indicate that invasive plants may regulate HFA effects by promoting the decomposition of native plants and increasing fauna abundance. Particularly, soil fauna groups play a very important role in this process. Our findings help us to re-understand the role of invasive plants in material cycling and energy flow in the context of achieving carbon neutrality goals.

1. Introduction

Litter decomposition is an essential ecological process that influences global carbon sink and material cycling [1,2,3]. The decomposition process is highly dependent on abiotic factors (e.g., mean temperature, precipitation, and moisture) on a global scale, and biological factors such as decomposer organisms, litter quality, and plant invasion on a local scale [2,4,5]. Many studies have proved that leaf litter tends to decompose more rapidly at a site dominated by the plant species from which it derived (i.e., home) compared to other sites (i.e., guest). This phenomenon is called the home-field advantage (HFA) effect [1,6,7]. In fact, the HFA effect was first used in sports competitions, and ecologists have conducted extensive research after introducing this concept [2,4]. The evidence of the HFA focused on specific decomposer communities that specialize in decomposing the litter of the home field, compared to litter from guest plant species [7,8,9]. Specialized soil fauna groups, the microbes, litter quality, and their interaction may have additive effects on the HFA during the litter decomposition process [10]. However, there is still limited research on the impact of invasive plant additions on the HFA effect during plant decomposition processes.
Species invasion is the primary problem facing seriously threatened ecosystems worldwide [11,12,13]. For instance, many invasive plants have caused ecosystem degradation by altering their species diversity, nutrient cycling, and decomposition process [14]. Invasive plants that can successfully invade are often characterized by a low C:N ratio [11,14]. Meanwhile, invasive plants may release leachates such as amino acids, monosaccharides, and polysaccharides during their decomposition phases [12,15]. These leachates are considered likely to promote the decomposition of more recalcitrant native plant [16,17]. The effect of invasive plants on the growth and decomposition of native plants has been well studied [5,11,13], but very little is known regarding the role of soil fauna groups in the HFA after the addition of invasive plants.
A large number of studies have shown the role of microbes in the HFA effect [7,8,9]. Specifically, the environmental conditions of the home field screen out microbial groups that can efficiently decompose litter [9]. These microbes have a certain preference for the types of litter in the home field and, in turn, these litters can provide sufficient nutritional sources for the microbes [8], thus forming a benign interaction [9]. Previous studies have suggested that soil fauna groups play a critical role in the HFA by directly or indirectly regulating the decomposition rate of litter [18,19,20]. Especially, soil macrofauna such as earthworms, earwigs, and isopods are essential drivers in transforming leaf litter into fine particulate organic matter in subtropical forests [21]. It is generally believed that the most efficient soil fauna groups are selected to decompose litter by the competition of litter resources [3,22]. Thus, soil fauna develops the specialized groups in degrading their litter [8,9]. The addition of invasive plants may regulate the specialization of soil fauna groups and activities of microbes [19], and then influence HFA [7,23].
Alternanthera philoxeroides (Mart.) Griseb. is a common amphibious invasive plant [24], which has a low C:N ratio and high concentrations of nutrients [24], and therefore is considered as high-quality litter. It can promote the decomposition of other species by releasing many leachates during their decomposition process [25]. These leachates may directly or indirectly regulate the activities of microbes and soil fauna groups, thus affecting the decomposition of litter at the home and guest sites [24,25]. To address the potential impact of the addition of invasive plants on the HFA during the litter decomposition process, we assembled litterbags with dry leaves of native Neosinocalamus affinis (Rendle) Keng f. (decomposition resistant) and Ficus virens W.T. Aiton (more easily decomposable) in single and double species mixtures with and without A. philoxeroides, and incubated these litterbags at N. affinis and F. virens sites at the edge of the forest for 360 days. We hypothesized that: (1) two native plants would exhibit the HFA effect during decomposition process; (2) the addition of A. philoxeroides would alter the magnitude of the HFA; (3) the addition of A. philoxeroides would increase the abundance of associated soil fauna groups.

2. Materials and Methods

2.1. Study Area

The study was conducted at the edge of a forest (107.43 E, 29.85 N; 195 m above sea level), situated on the banks of the Yangtze River in Chongqing city, China. The whole riparian zone is about 1000 m2. The surface soil in this area is mainly brown soil, and the deep soil is mainly limestone soil. During the 1-year study period, mean annual precipitation was 1094 mm and mean annual temperature was 18.5 °C. The forest age here is more than three decades. The plant communities involved are native Neosinocalamus affinis, Ficus virens, Cinnamomum camphora (L.) J. Presl and Cryptomeria japonica (Thunb. ex L. f.) D. Don, and invasive Alternanthera philoxeroides. A. philoxeroides is widely distributed at the edge of the forest. Resistant decomposable N. affinis litter and more easily decomposable F. virens litter are widely distributed in Chongqing city, and their tree layer coverage is more than 90%. After detachment, fallen leaves are often mixed with A. philoxeroides growing on the ground.

2.2. Experimental Design, Litterbag Preparation and Sampling

To test the HFA effect of riparian plant decomposition, two sites dominated by N. affinis and F. virens were selected for litter transplant experiment. The distance between the two sites was at least 300 m; the area of each site was about 100 m2. In each site, 18 randomized plots of 1 m × 1 m in a circular pattern were established with 10 m between two adjacent plots for litterbag incubation. There were four treatments (pure N. affinis, pure F. virens, N. affinis with A. philoxeroides, F. virens without A. philoxeroides) in each plot (18 plots × two sites × four treatments). The senescent leaves were collected by shaking the trees in autumn 2019. Sufficient A. philoxeroides (including roots, stems and leaves) were collected at the forest edge. The A. philoxeroides thalli were subsequently cut into sections (6–8 cm in length), like the length of N. affinis and F. virens leaves; all kinds of plant material were air dried for 1 month. After homogenizing the materials, the leaves of N. affinis and F. virens were placed in litterbags with and without A. philoxeroides. Litterbags (15 cm × 15 cm) were constructed from 5 mm nylon mesh, allowing access to soil fauna groups and microbes [21]. Each litterbag contained 6 g ± 0.1 of dry materials, and all species in any one combination were equally represented in mass.

2.3. Soil Sampling and Analysis

Top soil (10 cm) was sampled at each replicate (at least 50 m apart from each other) to measure the soil temperature, moisture, total C, total N, and pH at the beginning of the experiment and at six sampling times. Specifically, the soil temperature and pH were determined immediately in the field using the soil multiparameter probe (Fangke, FK-WSYP, Shandong, China) at each site to a depth of 10 cm. Before analysis, soil samples were stored in a refrigerator (Haier, BCD-572WDENU1, Qingdao, China) at 4 °C. Moisture was measured using a volumetric ring (Volume 100 mm3), oven drying at 105 °C for 24 h (DP, Y101A-1, Changzhou, China), and a hydrometer following the standard protocols [26]. Total C and total N were analyzed using the manual method of Bärlocher [27].
In total, 144 litterbags were randomly deposited at two sites on 1 July 2020 (two sites × four treatments × six sampling times × three replications = 144 litterbags). Litterbags were randomly deposited (pure N. affinis, pure F. virens, N. affinis with A. philoxeroides, F. virens without A. philoxeroides) in each plot. Dead branches and fallen leaves were removed from the surface of the forest floor, and plastic strings fixed the litterbags in position to prevent movement [26].

2.4. Taxa Measurement and Chemical Analyses

After 60, 120, 180, 240, 300 and 360 days, 36 litterbags at each sampling time were gently retrieved from randomly chosen trees at each site. Sampling time generally corresponded to the change of soil temperature within 1 year. Retrieved litterbags were immediately put into a cooler (JD, CL30, Chongqing, China), and returned to the laboratory to determine the mass remaining, faunal communities, and microbial respiration rate. The remaining leaf litter mass was expressed as the percentage of the initial mass.
In the laboratory, colonized macrofauna in the litterbags were picked out directly using tweezers and were fixed in 70% alcohol and identified to the genus or species level according to Yin et al. [28]. After homogenizing and weighing the wet litter in each litterbag, after removing excess water, a 0.5–3 g (depending on available material) subsample was used to estimate the wet mass–dry mass conversion factor to determine the dry mass of the entire sample. The abundance and richness of microfauna and mesofauna were quantified using 2 g of wet materials. They were transferred to 100 mL sample vials and 50 mL of sterile tap water was added. Subsamples were fixed using Lugol’s iodine solution and examined after 48 h using a microscope. After homogenizing the samples, 5 mL aliquots were removed using an autopipette and placed on a Perspex counting tray on a movable stage. The subsamples were evaluated under a stereomicroscope (Olympus SZ60, Tokyo, Japan) at a magnification of 40 × 10 until at least 300 individuals had been counted [19]. Taxonomic guides [28,29] were used to identify organisms at the genus or species level. Abundance of associated soil fauna groups was quantified as number of invertebrates/leaf dry mass (g).
The microbial respiration rate was quantified using 3 g of wet materials that remained in each litterbag according to Frouz et al. [30], which is a reliable indicator of the decomposition rate [31]. Specifically, the wet material (5 g) that remained in each litterbag was used to quantify microbial respiration rate, as a measure of overall microbial activity 31. Microbial respiration rates were measured as the amount of CO2 released from the samples without excess water. The leaf material was incubated in 500-mL airtight jars for 48 h at 25 °C in total darkness. The released CO2 was absorbed by 0.5 M NaOH solutions, and the amount of CO2 was determined by two-phase titration with 0.1 M HCl [32].
The initial litter qualities of A. philoxeroides, N. affinis and F. virens were determined by colorimetric analysis. The materials collected from the field were oven-dried at 40 °C for 72 h, and then they were ground to a fine powder for later use. Initial C, N and P concentrations were determined using a colorimetric analysis. The C concentration was analyzed using the H2SO4-K2Cr2O7 heat method, and the N and P concentration using colorimetric analysis [27]. The initial fiber concentration was extracted from the materials with neutral detergent and acid detergent (72% H2SO4) successively, and then ashed at 500 °C in a muffle furnace. The initial total phenol concentration was calculated using the Folin–Ciocalteau reagent employed by Bärlocher [27]. Specifically, after centrifuging the mixture of plant powder and acetone, the supernatant is absorbed and mixed with Folin Ciocalteau reagent and Na2CO3 solution to determine the total phenolic concentration through colorimetric analysis.

2.5. Data Analysis

The average decomposition rates (k) were calculated using the exponential decay model (Wt = W0 e−kt), where Wt is the dry litter mass remaining at day 360 and W0 is the dry litter mass at the initial time [27].
The HFA of the four treatments was also determined by calculating the mean HFA (% increase in k value at home compared with the guest site) for each litter type separately, following Austin et al. [9]:
M e a n   H F A ( % ) = ( k h o m e − k g u e s t ) / k g u e s t × 100
where, khome and kguest represent the decomposition constant of a given species at the home and at the guest site, respectively.
We calculated the HFA index (HFAI) to determine the overall HFA effects following Ayres et al. [8]. First, the relative mass loss was calculated as:
A R M L a ( % ) = A a A a + B a × 100
ARMLa is the relative mass loss of the litter from species A at site a, and Aa and Ba is the percentage (of initial) litter mass loss of plant species A and B decomposing at site a.
The HFA index (HFAI) was then calculated as:
H F A I % = A R M L a + B R M L b A R M L b + B R M L a × 100 − 100
where HFAI is the percent enhanced mass loss of litter when it decomposes at the home site versus the guest site and is a net value for both species (A and B) in the reciprocal transplant. HFAI for each sampling time was calculated to determine temporal changes in HFA effects.
The soil properties and initial chemical properties of three species, decomposition rates (k), mean HFA, and HFAI were compared using the Mann–Whitney U test and Kruskal–Wallis Test. After Log transformation (faunal abundance and microbial respiration rate) to meet the assumptions of normality and homoscedasticity, treatment effects on the responses were evaluated using three-way analysis of covariance (ANOVA) with sites, invasive plant addition and sampling time set as fixed factors, and mass remaining, faunal abundance and microbial respiration rate as dependent variables. Pearson correlation coefficients (r) were used to examine the correlation between decomposition rate and other factors (i.e., mean HFA, HFAI, the abundance of invertebrates and the microbial respiration rate). The significance threshold was set at p < 0.05. SPSS software (ver. 20, IBM, Armonk, NY, USA) was used to perform statistical analysis, and SigmaPlot 2000 (ver. 10, Systat Software, San Jose, CA, USA) was used to create Figures.

3. Results

3.1. Initial Litter and Soil Properties

Soil properties and initial litter qualities differed between the N. affinis and F. virens sites (Table 1 and Table 2). Soils at the N. affinis site had significantly higher total C concentrations but significantly lower moisture and total N concentrations than at the F. virens site (Table 1). Meanwhile, the initial qualities of A. philoxeroides had the highest N and P concentrations. N. affinis showed the highest C:N and C:P ratio. Ficus virens had the highest total phenol concentrations (Table 2).

3.2. Litter Decomposition and HFA

The remaining mass showed a downward tendency both at the N. affinis and F. virens sites (Figure 1), and it was significantly affected by the sites (F1,80 = 30.70, p < 10−4), addition of invasive plants (F1,80 = 174.93, p < 10−4) and sampling time (F5,80 = 420.36, p < 10−4; Table 3). The decomposition rate of litter at the home site was always significantly higher than that at the guest site whether invasive plants were added or not (Table 4). Furthermore, their decomposition rates increased significantly after adding the invasive A. philoxeroides (Table 4).
The four treatments all showed a positive HFA effect (Figure 2). Among them, the mean HFA of N. affinis litter was significantly higher than that of other three treatments (Figure 2). Positive HFA effects with litter mass loss were at least 1.32% faster at home than at the guest sites. The mean HFA effect was altered by adding A. philoxeroides in two species (Figure 2). The addition of A. philoxeroides reduced the mean HFA of N. affinis litter and increased that of F. virens litter (Figure 2). The HFAI of litter decomposition for the two species with and without A. philoxeroides were all positive (Figure 3). The HFAI without A. philoxeroides was significantly higher than that with A. philoxeroides (F1,34 = 4.68, p = 0.038; Figure 3).

3.3. Leaf-Associated Faunal Community and Microbial Respiration Rate

During the experiment, 36 soil fauna groups were recorded. They were dominated by the protist group Sarcodina (16 taxa), followed by the Arthropod group Arachnida (3 taxa; see Supplementary Table S1). Their abundance in the colonized litterbags at the home site was always higher than at the guest site (Figure 4). Furthermore, the addition of A. philoxeroides increased the abundance of associated soil fauna groups in two species. Furthermore, the abundance of springtails (F1,14 = 13.03, p = 0.003), spiders (F1,22 = 5.77, p = 0.025), ants (F1,14 = 9.66, p = 0.008) and silverfishes (F1,6 = 11.87, p = 0.014) at the N. affinis site was significantly higher than that at the F. virens site, and that of woodlouse (F1,14 = 6.05, p = 0.028), earthworms (F1,14 = 61.18, p < 10−4), nematodes (F1,14 = 18.29, p = 0.001) and earwigs (F1,14 = 10.09, p = 0.007) at the N. affinis site was significantly lower than that of the F. virens site (Table S1). Their abundance was significantly affected by the sites (F1,80 = 72.41, p < 10−4), addition of invasive plants (F1,80 = 11.15, p = 0.001), sampling time (F5,80 = 270.35, p < 10−4) and the interaction of the site and sampling time (F5,80 = 3.46, p = 0.006; Table 3).
The microbial respiration rates showed a tendency of first increasing and then decreasing, and the peak was at day 120 at two sites (Figure 5). No significant difference was observed between the N. affinis and F. virens sites (F1,142 = 0.07, p = 0.794). The microbial respiration rate was significantly affected by the sampling time (F5,80 = 23.59, p < 10−4) and the interaction of the addition of invasive plants and sampling time (F5,80 = 5.41, p < 10−4; Table 3).
Pearson correlation coefficients (r) showed that the decomposition rate was significantly correlated with HFAI at the two sites, and with the invertebrate abundance at the N. affinis site. There was no significant correlation between the decomposition rate and microbial respiration rate at the two sites (Table 5).

4. Discussion

4.1. The Impact of Initial Quality and Environmental Conditions on HFA

Soil surface receives large inputs of fallen leaves, and their initial quality is one of the predominant drivers of the HFA [4]. In the present study, a positive HFA effect was presented at two selected sites. The main reasons may be attributed to the different initial qualities of litter and the specialization of soil fauna groups and microbes in degrading their litter [4,8]. In nature, plant litter with high initial quality tends to have low initial C:N, C:P, and N:P ratios [33], which affect the litter mass remaining and the direction and strength of the HFA [9]. In the present study, the initial quality of A. philoxeroides was the highest, followed by F. virens and N. affinis. The mean HFA of N. affinis litter was significantly higher than that of other three treatments. The possible explanation is that the HFA often presents stronger effects on recalcitrant litter compared to fast-decomposing litter 8. In addition, the litter of F. virens has higher N and P concentrations, which was considered to promote colonization by microbes, and then increase colonization by specialized fauna groups [17]; this may be the reason why the mean HFA in the treatment of F. virens was lower.
The site-specific environmental conditions strongly affect the interactions between plant litter and soil environments for plant decomposition [34], and explain the occurrence and magnitude of the HFA largely by mediating the activity of soil fauna groups and microbial communities [35,36]. In our study, the initial leaves of N. affinis contains higher C and lower P concentrations than that of F. virens. Compared with the F. virens site, significantly higher C and lower N concentrations at the N. affinis site were observed. This result is consistent with a previous study, which revealed that higher C concentrations in litter and organic matter concentrations in soil could enhance HFA, yet higher P concentrations in litter and N concentrations in soil can weaken HFA [37]. The most likely reason is that litter with high carbon concentrations has developed specialized fauna groups and microbes at the home site, and these specialized decomposers are difficult to develop in a short time away from home, thus higher C concentrations in litter can promote the HFA effect [9]. In other words, highly efficient decomposition microbial and fauna community were screened from high carbon litter [35,36]. Once the concentrations of N and P in the soil increases, it may provide additional resources for soil microbes, greatly shortening the time for the development of specialized decomposers, thus weakening the HFA effect [37,38].
There was no significant difference in soil temperature between two selected sites, but a significant difference was observed in moisture. Therefore, it can be conjectured that the difference in the decomposition rate may be driven by moisture on a local scale [21], and the potential increase in decomposition rate for the N. affinis site with temperature appears to be mitigated by the lower soil moisture [39]. Soil moisture may affect plant decomposition by directly affecting leaching processes [40] and indirectly affecting the distribution of soil fauna groups and microbial activity [41,42].

4.2. HFA Effects in Response to the Addition of A. philoxeroides

Labile and fast-decomposing litter is expected to stimulate decomposer activity [16], and then affect the magnitude of the HFA [3]. To our knowledge, few studies have investigated how the addition of invasive plants influences HFA. It has been reported that nitrogen enrichment caused by artificial fertilization could lead to smaller HFA effects through altering microbial abundance and community composition [26]. Our experimental results showed that the addition of A. philoxeroides reduced the mean HFA of N. affinis litter and increased that of F. virens litter. HFAI without A. philoxeroides was significantly higher than that with A. philoxeroides. Pearson correlation coefficients (r) showed that the decomposition rate was significantly negative correlated with F. virens litter at the two sites. One possible explanation is that leachates produced by adding A. philoxeroides may have increased low-quality litter turnover at the home and guest sites. The addition of A. philoxeroides in N. affinis litter can stimulate the activity of soil fauna groups and microbial communities [13] and composition, and allow nutrient transfer to the low-quality litter [41], which therefore loses mass more rapidly and decomposes faster [13], thus reducing the HFA effect. Meanwhile, this stimulation is not coupled with an increased F. virens turnover because microbes likely allocated leachates to reproduction, growth or respiration instead of C mineralization [17]. Additionally, F. virens is rich in higher total phenols, which are believed to inhibit microbial activity [43,44,45].
The co-occurrence of A. philoxeroides and leaf litter at the edge of the forest is a necessary condition for present study [46]. Changing the quality of leaf litter may alter the magnitude and direction of the HFA due to decomposer colonization [3,17]. As predicted by our second hypothesis, the addition of invasive plants accelerated the decomposition rate yet altered the mean HFA. This implies that a specialized soil community driving the HFA can accelerate litter decomposition [3]. The HFA tends to present a stronger effect on low-quality litter compared to high-quality litter [3], and depends on the invaded ecological environment [9].

4.3. The Role of Soil Fauna Groups and Microbes

The soil microbial community has received more attention in the HFA [7,9,34]. In recent years, the importance of soil fauna groups to ecosystem functions has also been highlighted [21,30]. According to Ayres et al. [8], the HFA can be considered as a phenomenon in which different quality litter may select the most efficient soil fauna groups to decompose their litter matrix by competing as a source of energy and nutrients. In our experiment, faunal abundance colonizing the home site was always higher than that at the guest site. Collembola (springtails), Arachnida (spiders and mites) and Formicidae (ants) preferred colonization at the N. affinis site, and more Isopoda (woodlouse), Oligochaeta (earthworms), nematodes and Dermaptera (earwigs) were observed at the F. virens site. The most likely explanation is that fungi colonize the leaves of N. affinis with higher cellulose concentrations, which provides rich food resources for springtails and ants, and attracts more predators (spiders) to colonize there [47]. Their foraging and burrowing activities largely influence the physical environment they live in, consequently promoting nutrient flow and material cycling [21]. Simultaneously, the soil fauna groups at the F. virens site may prefer a wetter environment with higher nutrient concentrations. For instant, earthworms have been described as ecosystem engineers, increasing the mineralization and decomposition of organic matter through direct feeding on litter and activity [48]. Nematodes and woodlice make significant contributions to litter decomposition, and their abundance was positively correlated with soil moisture and negatively correlated with soil temperature [21]. The soil fauna living at the home site for a long time have better adapted to their surroundings, thus developing specialized soil fauna groups [8,23]. Furthermore, the addition of high-quality invasive plants provides an additional source for these specialized soil fauna groups, which may make them more efficient in decomposing litter [19], thus becoming a key factor affecting the decomposition of litter [49]. Compared with the guest site, these specialized soil fauna groups are better at decomposing the litter at the home site, thus producing the HFA effect [10].
The presence of invasive plant litter may release leachates (e.g., amino acids, monosaccharides, and polysaccharides) during the decomposition process [12], which provide high-quality resource to microbial communities and indirectly increase the consumption of litter by fauna groups [50]. In turn, these soil fauna and microbes may be specialized in the decomposition of localized litter, which can lead to enhance mineralization [23] and regulate the HFA effect. In our experiment, Pearson correlation coefficients (r) showed that the decomposition rate was significantly positively correlated with the abundance of invertebrates colonizing F. virens litter at the two sites, but that of N. affinis was only significantly related at the home site. It seems that invertebrates at the N. affinis site are more specific. The most likely explanation is that it may involve the initial mass of litter and the soil characteristics of the site [4,8]. The experimental results showed that the fiber concentration of N. affinis was significantly higher than that of F. virens, and the soil moisture at the N. affinis site was significantly lower than that at the F. virens site. The springtails colonizing the N. affinis site seemed to prefer colonization in a drier environment and fed on fungi colonizing the fiber, and the springtails were eaten by spiders and ants [51].
In the forest, macrofauna can convert leaf litter into small particles without directly participating in the mineralization of organic matter [21,30]. Meanwhile, they increase the activities of the soil micro- and mesofauna which are responsible for fine-particle decomposition [21]. Specifically, micro- and mesofauna can convert fine particle matter into ultrafine particles through their activities [52,53]. Furthermore, their activities promote microbial population renewal by predating senescent microbial cells [54]. Increasing the activity of microbial populations in turn promoted recycled nutrients [36]. The surface area of the leaf substrate is increased by the fragmentation of macrofauna activity. It allows more microbes to colonize the internal tissues of litter [53], which makes the litter produce fine particles that can potentially be used by micro- and meiofauna [52].

5. Conclusions

In this study, we observed that the presence of the invasive plant A. philoxeroides could alter the home-field advantage of native leaf litter decomposition. After adding invasive plants, the mean HFA of N. affinis litter decreased significantly, while that of F. virens litter increased slightly. In addition, the addition of invasive plants also significantly reduced HFAI. This could be attributable to initial leaf quality and taxonomic and functional diversity of associated soil fauna groups in different site environments. We further emphasized the regulatory role of initial litter quality and soil fauna groups on the interaction between the invasive effect and the HFA. Moreover, we found a significant difference in faunal abundance between N. affinis and F. virens sites, implying that soil fauna groups would have environmental specialization. Our findings show that the addition of an invasive plant regulates the home-field advantage, and highlights the important role of soil fauna groups in this process. Given the widespread invasion of A. philoxeroides all over the world, our research indicates that native plant decomposition regulated by A. philoxeroides will help us with forest carbon budgeting.

Supplementary Materials

The following supporting information can be downloaded at: https://www.mdpi.com/article/10.3390/f15101708/s1; Table S1: The mean faunal abundance (number bag-1) according to two sites of the litterbags with and without of A. philoxeroides over 360 days (n = 3).

Author Contributions

Methodology, S.C. and H.Z.; Software, S.C., X.X., J.W., H.W., Y.J. and Z.G.; Investigation, S.C., H.Z., H.W., Y.J. and Z.G.; Writing—original draft, S.C. and X.X.; Writing—review & editing, S.C. and J.W.; Funding acquisition, S.C. and J.W. All authors have read and agreed to the published version of the manuscript.

Funding

This study was supported by grants from the National Key Research and Development Project (2022YF3201704); Scientific and Technological Research Program of Chongqing Municipal Education Commission [KJQN202201428].

Data Availability Statement

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

Conflicts of Interest

The authors declare that they have no conflicts of interests.

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Forests 15 01708 g001
Figure 1. Remaining mass of Neosinocalamus affinis, Ficus virens, and mixed with or without invasive Alternanthera philoxeroides at the Neosinocalamus affinis and Ficus virens sites. Data are the mean ± SE.
Figure 1. Remaining mass of Neosinocalamus affinis, Ficus virens, and mixed with or without invasive Alternanthera philoxeroides at the Neosinocalamus affinis and Ficus virens sites. Data are the mean ± SE.
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Figure 2. The mean home-field advantage (mean HFA) for Neosinocalamus affinis, Ficus virens, and with or without invasive Alternanthera philoxeroides. Data are the mean ± SE. Different lowercase letters indicate the significance among different treatments of mean HFA (p < 0.05).
Figure 2. The mean home-field advantage (mean HFA) for Neosinocalamus affinis, Ficus virens, and with or without invasive Alternanthera philoxeroides. Data are the mean ± SE. Different lowercase letters indicate the significance among different treatments of mean HFA (p < 0.05).
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Figure 3. Parameter estimates calculated for the home-field advantage index (HFAI). Data are the mean ± SE.
Figure 3. Parameter estimates calculated for the home-field advantage index (HFAI). Data are the mean ± SE.
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Figure 4. Abundance of associated soil fauna groups in decomposing leaves of Neosinocalamus affinis, Ficus virens, and mixed materials with or without invasive Alternanthera philoxeroides at the Neosinocalamus affinis and Ficus virens sites. Data are the mean ± SE.
Figure 4. Abundance of associated soil fauna groups in decomposing leaves of Neosinocalamus affinis, Ficus virens, and mixed materials with or without invasive Alternanthera philoxeroides at the Neosinocalamus affinis and Ficus virens sites. Data are the mean ± SE.
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Figure 5. Microbial respiration rates according to the litterbags with or without Alternanthera philoxeroides at the Neosinocalamus affinis and Ficus virens sites. Data are the mean ± SE.
Figure 5. Microbial respiration rates according to the litterbags with or without Alternanthera philoxeroides at the Neosinocalamus affinis and Ficus virens sites. Data are the mean ± SE.
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Table 1. Chemical and physical properties of soils at the Neosinocalamus affinis site and the Ficus virens site.
Table 1. Chemical and physical properties of soils at the Neosinocalamus affinis site and the Ficus virens site.
SitesSoil Temperature (°C)Soil Moisture (%)Total C (g Kg−1)Total N (g Kg−1)pH
N. affinis17.27 ± 0.072 ns21.04 ± 0.941 *52.43 ± 1.793 *3.38 ± 0.032 **6.86 ± 1.032 ns
F. virens14.03 ± 0.08923.96 ± 0.49247.34 ± 1.8395.54 ± 0.0486.35 ± 0.932
Notes: Values are presented as mean ± SE (n = 3). Mean values were compared by Mann–Whitney U test. Within columns, significant differences between habitats are given by: * p < 0.05; ** p < 0.01, ns p > 0.05.
Table 2. Initial chemical properties of Alternanthera philoxeroides, Neosinocalamus affinis, and Ficus virens litter.
Table 2. Initial chemical properties of Alternanthera philoxeroides, Neosinocalamus affinis, and Ficus virens litter.
FactorsA. philoxeroidesN. affinisF. virens
C (% DM)11.95 ± 0.360 a41.67 ± 0.001 b25.77 ± 0.069 c
N (% DM)4.21 ± 0.012 a0.48 ± 0.024 b1.66 ± 0.052 c
P (% DM)2.29 ± 0.076 a0.05 ± 0.034 b0.14 ± 0.076 c
C: N3.75 ± 0.025 a86.79 ± 0.790 b15.55 ± 0.025 c
C:P6.87 ± 0.065 a833.21 ± 0.973 b184.46 ± 0.065 c
N:P1.81 ± 0.022 a9.57 ± 0.007 b11.86 ± 0.022 b
Total phenol (% DM)5.43 ± 0.091 a1.40 ± 0.037 b10.10 ± 0.091 c
Lignin (% DM)2.67 ± 0.003 a3.16 ± 0.002 b8.52 ± 0.003 b
Cellulose (% DM)3.21 ± 0.008 a25.28 ± 1.566 b17.08 ± 0.008 b
Hemicellulose (% DM)8.97 ± 0.059 a49.72 ± 2.893 b12.78 ± 0.059 c
Notes: Values are presented as mean ± SE (n = 3). Mean values were compared using the Kruskal–Wallis Test. Significant differences in initial quality factors between the three species are given by different superscript letters. DM, dry mass.
Table 3. Results of three-factor analysis of variance (ANOVA): site (S), addition of invasive plant (A), sampling time (T), and their interactions on the mass remaining, faunal abundance, and microbial respiration rates over 360 days.
Table 3. Results of three-factor analysis of variance (ANOVA): site (S), addition of invasive plant (A), sampling time (T), and their interactions on the mass remaining, faunal abundance, and microbial respiration rates over 360 days.
SourcedfMass RemainingFaunal AbundanceMicrobial Respiration Rate
FpFpFp
S130.70<10−472.41<10−40.150.704
A1174.93<10−411.150.0010.160.692
T5420.36<10−4270.35<10−423.59<10−4
T × A11.740.1900.400.5300.160.692
T × S50.230.9483.460.0060.690.631
A × S50.450.8130.840.5255.41<10−4
S × A × T50.080.9950.060.9980.340.889
Table 4. Comparison of the decomposition rates according to the litterbags placed at Neosinocalamus affinis site and Ficus virens site over 360 days.
Table 4. Comparison of the decomposition rates according to the litterbags placed at Neosinocalamus affinis site and Ficus virens site over 360 days.
SitesDecomposition Rate (k·Day−1)
N. affinisF. virensN. affinis × A. philoxeroidesF. virens × A. philoxeroides
N. affinis site0.003 ± 0.001 a,*0.002 ± 0.001 b,*0.004 ± 0.001 c,*0.003 ± 0.001 a,*
F. virens site0.002 ± 0.001 a0.003 ± 0.001 b0.003 ± 0.001 b0.004 ± 0.001 c
Notes: Data are the mean ± SE. Mean values were compared by the Mann–Whitney U test and Kruskal–Wallis Test. Different lowercase superscript letters indicate the significance of different materials at the same sites (p < 0.05). Within columns, significant differences between sites are given by: * p < 0.05.
Table 5. Pearson correlation coefficients (r) between decomposition rate and other factors.
Table 5. Pearson correlation coefficients (r) between decomposition rate and other factors.
FactorsSiteN. affinisF. virensN. affinis × A. philoxeroidesF. virens × A. philoxeroides
rprprprp
Mean HFAN. affinis site−0.6520.161−0.8420.035−0.0140.979−0.6550.158
F. virens site−0.6770.139−0.8170.047−0.0430.935−0.6060.202
HFAIN. affinis site−0.8920.017−0.8810.020−0.8900.017−0.8950.016
F. virens site−0.9010.014−0.8700.024−0.8780.021−0.8650.026
Invertebrate abundanceN. affinis site0.8700.0240.8880.0180.8150.0480.8730.023
F. virens site0.7930.0600.8520.0310.7900.0620.8890.018
Microbial respiration rateN. affinis site−0.0950.857−0.0590.911−0.0980.854−0.0290.956
F. virens site−0.0630.906−0.1270.811−0.0880.869−0.0660.901
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Chen, S.; Xie, X.; Wen, J.; Zhai, H.; Wang, H.; Jiang, Y.; Gou, Z. The Addition of an Invasive Plant Alters the Home-Field Advantage of Native Leaf Litter Decomposition. Forests 2024, 15, 1708. https://doi.org/10.3390/f15101708

AMA Style

Chen S, Xie X, Wen J, Zhai H, Wang H, Jiang Y, Gou Z. The Addition of an Invasive Plant Alters the Home-Field Advantage of Native Leaf Litter Decomposition. Forests. 2024; 15(10):1708. https://doi.org/10.3390/f15101708

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Chen, Shaojun, Xiaohua Xie, Jie Wen, Hao Zhai, Huiqi Wang, Yuhang Jiang, and Zhanxu Gou. 2024. "The Addition of an Invasive Plant Alters the Home-Field Advantage of Native Leaf Litter Decomposition" Forests 15, no. 10: 1708. https://doi.org/10.3390/f15101708

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