Litter quality drives the shift of soil fauna structure without changing its ecological function

Litter quality drives the shift of soil fauna structure without changing its ecological function | Research Square window.SnipcartSettings = { analytics: { enabled: false } }; (function() { var accessVector = localStorage.getItem('access_vector') || ''; window.dataLayer = window.dataLayer || []; if (accessVector) { window.dataLayer.push({ user: { profile: { profileInfo: { snid: accessVector } } } }); } })(); (function(w,d,s,l,i){w[l]=w[l]||[];w[l].push({'gtm.start':new Date().getTime(),event:'gtm.js'});var f=d.getElementsByTagName(s)[0],j=d.createElement(s),dl=l!='dataLayer'?'&l='+l:'';j.async=true;j.src='https://www.googletagmanager.com/gtm.js?id='+i+dl;f.parentNode.insertBefore(j,f);})(window,document,'script','dataLayer','GTM-K279D39R'); Browse Preprints In Review Journals COVID-19 Preprints AJE Video Bytes Research Tools Research Promotion AJE Professional Editing AJE Rubriq About Preprint Platform In Review Editorial Policies Our Team Advisory Board Help Center Sign In Submit a Preprint Cite Share Download PDF Research Article Litter quality drives the shift of soil fauna structure without changing its ecological function Kui Long, Zhenyu Zhou, Gan Yu, Yakun Zhang, Yang Wang, Qingyun Wang, and 2 more This is a preprint; it has not been peer reviewed by a journal. https://doi.org/ 10.21203/rs.3.rs-6625409/v1 This work is licensed under a CC BY 4.0 License Status: Published Journal Publication published 03 Sep, 2025 Read the published version in Plant and Soil → Version 1 posted 6 You are reading this latest preprint version Abstract Background and aims Invasive plant species represent significant threats to ecosystems globally, disrupting native habitats and adversely affecting biodiversity and ecosystem functions and services. This study focused on the impact of bamboo invasion, specifically the spread of moso bamboo ( Phyllostachys edulis ), on soil fauna communities in subtropical forests. Methods Using litter reciprocal transplant experiments between uninvaded broadleaf forests and invaded bamboo forests, we investigated how alterations in litter quality influenced soil fauna diversity and community composition. Results Reciprocal litter transplants primarily affected soil fauna community composition through species replacement without changing their diversity or ecological function. Notably, bamboo litter, which is considered high quality, had a more pronounced impact on soil fauna communities in low-quality litter environments but not vice versa. Conclusion These findings highlight the intricate relationship between litter quality, soil fauna communities, and ecosystem stability in the context of plant invasion. Understanding these dynamics provides valuable insights for ecosystem management strategies in the face of invasive plant species. Plant invasion Litter reciprocal transplants Soil fauna communities Litter quality Feeding guilds Figures Figure 1 Figure 2 Figure 3 Figure 4 Figure 5 1. Introduction Invasive alien plant species are recognized as driving biodiversity loss and altering ecosystem services and socioeconomic conditions (Bacher et al., 2018 ; Rai and Singh, 2020 ; Pérez et al., 2022 ; Gallardo et al., 2024 ). The ecological disturbances caused by invasive plants have been increasingly recognized as a key challenge to global sustainability (Rai and Singh, 2020 ; Premakumari et al., 2022 ; Khattak et al., 2024 ). These species impede native plant growth by affecting soil cover, nutrient cycling, fire patterns, and hydrology (Weidlich et al., 2020 ; Raheem et al., 2024 ) and trigger a series of environmental effects that reshape community composition, biotic interactions (Vilà et al., 2011 ; David et al., 2017 ; Li et al., 2022 ; Zhang, 2023b ), and ecosystem processes, influencing ecosystem productivity and human well-being (Xu et al., 2020 ). In recent years, the invasion of moso bamboo ( Phyllostachys edulis ) has emerged as an ecological concern, not only in its native region in China but also in areas where it has been introduced, such as Japan and North America (Fukushima et al., 2015 ; Lieurance et al., 2018 ; Xu et al., 2020 ). As a native species in subtropical China, moso bamboo is the most prevalent giant timber bamboo species and is extensively cultivated (Peng et al., 2013 ; Li et al., 2024 ). However, in recent decades, abandoned bamboo plantations have led to the spread of moso bamboo into neighboring broadleaf forests, negatively affecting plant diversity, litter input, soil properties, and soil communities (Liu et al., 2019 ; Luan et al., 2021 ; Wei et al., 2021 ; Li et al., 2024 ). The impacts are primarily manifested as changes in litter input, root exudates, and the physical environment, facilitating invasion success by increasing primary productivity and affecting soil biota and ecosystem dynamics (Wolfe and Klironomos, 2005 ). Moso bamboo invasion changes the biodiversity and community of soil fauna (Wei et al., 2021 ; Long et al., 2023 ; Xiao et al., 2023 ). Litter from invasive plants often decompose faster than that from native plants, altering soil carbon storage and nutrient levels (Ehrenfeld, 2010 ). These changes in litter quantity and quality modify habitat properties, such as soil moisture and pH, affecting soil biota performance (Kappes et al., 2007 ; Zhao et al., 2021 ; Morris and Blackwood, 2024 ). Research has demonstrated that alterations in litter inputs from invasive plants significantly affect soil organisms, including the abundance and diversity of soil biota species, thereby influencing decomposition processes and nutrient cycling (Liao et al., 2008 ; Sun et al., 2023 ; Zhang, 2023a ). Previous studies have primarily focused on the impacts of microorganisms; thus, the specific effects of soil fauna remain uncertain. Soil fauna significantly impact decomposition processes and serve as important biodiversity reservoirs, while their diversity and functions vary and are closely linked to different aboveground systems (Peng et al., 2020 ; Wang et al., 2020 ; Coleman et al., 2024 ). However, there is no definitive conclusion regarding their correlation. Soil fauna are integral ecosystem components, playing crucial roles in accelerating organic matter decomposition and nutrient cycling, primarily by breaking down litter and stimulating microbial activity (Petersen, 1982 ; Seastedt and Crossley Jr, 1983 ; Bardgett and Chan, 1999 ; Yin et al., 2010 ; Reddy et al., 2024 ). Soil fauna from different functional groups contribute differently to litter decomposition, with the most fundamental functional distinctions being shredders (involved in fragmentation), i.e., detritivores, and predators (Faber and Verhoef, 1991 ; Brussaard, 1998 ; Heděnec et al., 2022 ; Coleman et al., 2024 ). Shredders break down plant litter, increasing food resources for soil microfauna and mesofauna (Brown, 1995 ; Olivia and Dumitru, 2023 ). Fragmentation enhances the surface area of litter substrates, allowing microbes to access nutrient-rich inner tissues and thereby influencing litter decomposition (Faber and Verhoef, 1991 ; Chapin et al., 2002 ; Coleman et al., 2017 ; Cavallet et al., 2022 ). As evidenced, changes in the soil fauna community composition impact litter decomposition and nutrient metabolism (Verhoef and Brussaard, 1990 ; Setälä et al., 1996 ; Jones et al., 1998 ; Li et al., 2021 ; Yang et al., 2022 ; Balasubramanian et al., 2023 ). Would changes in litter (e.g., litter quality) affect soil fauna? Peng et al. ( 2022 ) speculated that leaf litter quality likely serves as the primary driver for the effects of tree species on soil fauna communities. However, this speculation has not yet been confirmed. Litter transplant is an effective method of litter control that confirms the “home-field advantage” (HFA) hypothesis, which states that decomposition occurs more quickly when litter decomposes beneath the plant species from which it originates (i.e., at home) than beneath different plant species (away) (Bocock et al., 1960 ; Hunt et al., 1988 ; Gholz et al., 2000; Negrete-Yankelevich et al., 2008 ; Ayres et al., 2009a ). Previous studies have shown that litter transplant changes the microbial diversity and community structure (Milcu and Manning, 2011 ; Lin et al., 2019 ). Dynamic changes in microbial communities in litter are instrumental in elucidating changes in component composition and material cycling during decomposition (Ma et al., 2023 ). Therefore, researchers are interested in the effects of the decomposition process on microbial communities in litter transplant experiments (Zhong et al., 2018 ), while research on soil fauna is still relatively scarce. The reciprocal transplant of litter between invasive and native plants offers a potential strategy for investigating the effects of altering litter inputs on soil fauna communities. Understanding how alternations in litter quality during plant invasion affect decomposers, such as soil fauna that act as shredders, will provide further insights into ecosystem dynamics and management strategies. We focused on moso bamboo as the experimental subject to investigate the effects of litter quality on soil fauna in plant invasions. We conducted litter exchange and retention treatments between two forest stands, i.e., secondary broadleaf forest and Phyllostachys edulis forest, representing two invasion stages: uninvaded and completely invaded. We investigated the impact of bamboo invasion on soil fauna diversity at both the taxonomic and functional levels as well as their community composition. For this experiment, the following scientific questions were posed: (1) Does reciprocal litter transplant (exchanging litter of different qualities) affect soil fauna diversity and community structure? (2) Does changing litter quality through reciprocal transplant affect the ecological functions performed by soil fauna? 2. Materials and methods 2.1 Study site The study site was in the Tianmu Mountain National Nature Reserve, located in Zhejiang Province, Eastern China (30°18'–30°25'N, 119°23'–119°29'E) at an elevation range of 500–650 m. This region experiences a subtropical monsoon climate with an average annual temperature of 8.8–14.8°C and an average annual rainfall of 1390–1870 mm. The soil in the study site was derived from silt stone, classified as a Ferralsol according to the Food and Agriculture Organization of the United Nations soil classification system (WRB, 2006 ). Following the abandonment of bamboo forests in the 1970s, the natural forest at this site heavily affected moso bamboo ( Phyllostachys edulis ) encroachment (Chen et al., 2021 ). At this site, we identified three distinct forest types, each representing a different stage of bamboo invasion (Wei et al., 2021 )): secondary broadleaf forest (SBF), an uninvaded native forest dominated by Cunninghamia lanceolata , Quercus serrata var. brevipetiolata , and Pinus massoniana ; mixed bamboo forest (MBF), a transition zone moderately invaded by moso bamboo; and Phyllostachys edulis forest (PEF), which is completely invaded and dominated by moso bamboo. Our research selected SBF (uninvaded) and PEF (completely invaded) at the site for the present study. Both had similar elevations, slopes (50–60%), soil textures, and bulk densities (Teng et al., 2023 ). 2.2 Experimental design Six parallel transects (about 180 m in length and 50 m apart) along the encroaching path from broadleaf forest to moso bamboo forest were installed in 2016 in an area of about 4.5 ha (Wei et al., 2021 ). For this study, four paired plots (20 m × 20 m) of broadleaf forest and moso bamboo forest in the six transects were randomly selected (for four replicates). In each plot, three subplots (each 2.5 m × 2.5 m, 10–15 m apart) were established (for three treatments), resulting in 24 subplots (Fig. S1 ). Nylon net fences, standing 0.3 m high with a mesh size of 5 mm, were installed around the perimeter of each subplot to prevent intrusion of the surrounding litterfall. Additionally, a top nylon net (2.5 m × 2.5 m, 2 mm mesh size) was suspended 0.8 m above each subplot to intercept fresh litterfall falling into the plot (Fig. S2). These subplots were set up in February 2019. All aboveground litter within each subplot was cleared, followed by a 3-month buffer period. These three subplots in each plot were established in May 2019 and arranged randomly for three monthly treatments of the intercepted fresh litterfall: a) control group: litterfall retention—fresh litterfall intercepted by the top nets from each forest stand was mixed monthly and evenly redistributed onto their own subplot; b) removal group: litterfall removal—fresh intercepted litterfall was cleared monthly; and c) replacement group: litterfall replacement—fresh litterfall intercepted from broadleaf or bamboo forests was exchanged and scattered onto the subplots of the other forest type (Fig. S1 ). The litterfall in each litter trap was collected monthly and evenly added to the control and replacement treatments at an ambient rate. The average rate of litter input was 0.3 kg dry litter per m 2 per year for broadleaf forests and 0.33 kg dry litter for bamboo forests (Teng et al., 2023 ). 2.3 Soil fauna extraction In May 2022 (3 years after litterfall manipulation), November 2022 (3.5 years), and May 2023 (4 years), soil fauna diversity was investigated in each subplot. Litter and soil samples were collected using the three-point sampling method in each subplot. For each point, litter and humus from an area of 25 cm × 25 cm as well as a soil column with a diameter of 5 cm and a depth of 20 cm were collected. The collected samples were labeled and transported to the laboratory on the same day. The soil fauna were then extracted using the Berlese–Tullgren funnel method (2-mm diameter mesh) (Macfadyen, 1953 ) in 48 h and gathered in conical flasks containing anhydrous ethanol. Soil fauna were preliminarily classified and categorized based on their morphology, as observed under a microscope (VHX-5000, Keyence Corporation, Osaka City, Japan). In each plot, soil fauna individuals from litter or soil samples from the three sampling points were bulked into one composite sample. In total, 120 samples were collected, with 72 from the soil layer (2 forest types × 3 litter manipulations × 4 replicates × 3 times sampling) and 48 from the litter layer (no samples from the removal treatment). The three collected samples were considered as repeated measurements. The litterfall from the removal subplots in the two forest types was collected in November and December 2022. Since the two months encompassed the peak leaf-fall period for both bamboo and broadleaf forests, the litterfall was used to determine the initial physicochemical properties of litter inputs (the contents of cellulose, lignin, total carbon, and total nitrogen), following the protocols described by Teng et al. ( 2023 ) (Table S1 ). 2.4 High-throughput sequencing (HTS) and data processing We employed a highly efficient and reliable high-throughput sequencing (HTS) technique (Arribas et al., 2016 ; Creedy et al., 2019 ) to acquire the DNA sequences of the soil fauna (Wei et al., 2021 ; Long et al., 2023 ). All 120 samples underwent grinding and homogenization with liquid nitrogen for DNA extraction using the DNeasy® Blood and Tissue Kit (QIAGEN, Hilden, Germany) following the sequencing and data processing protocols outlined by Wei et al. ( 2021 ). Primers Ill_B_F (5′. CCIGAYATRGCITTYCCICG. 3′) (Shokralla et al., 2015 ) and Fol_degen_rev (5′. TANACYTCNGGRTGNCCRAARAAYCA. 3′) (Yu et al., 2012 ) were utilized in polymerase chain reaction (PCR) to amplify the 418-bp region of the cytochrome oxidase subunit I (COI) gene (barcode fragment). Each sample was amplified in triplicate on an ABI GeneAmp®9700 (Applied Biosystems, Waltham, Massachusetts, USA), following the PCR protocol outlined by Arribas et al. ( 2016 ). After the dual-index barcodes (Illumina TruSeqTM DNA Sample Prep Kit, Illumina, Inc., San Diego, California, USA) were added, the purified amplicons were pooled and sequenced on the Illumina MiSeq Platform (2 × 300 bp paired-end) following quality control. The raw sequences underwent quality filtration using Trimmomatic (Bolger et al., 2014 ) and were merged using FLASH (Magoč and Salzberg, 2011 ). Only contigs with a length of 418 bp were retained. Post-joining, the DNA sequences were processed and optimized using the DADA2 denoising method to obtain amplicon sequence variants (ASVs). Subsequently, TaxonDNA software (version 1.0) was employed to cluster the ASVs at a 97% threshold to generate operational taxonomic units (OTUs). To prevent sequencing errors, we removed low-abundance OTUs (total counts < 5). Most soil fauna sequences generated from field biodiversity investigations could not be identified to the species level from any reference database, lacking deep coverage for soil invertebrates (Wei et al., 2021 ; Long et al., 2023 ). The data sorting procedure in this study fully combined the broad advantages of the two dominant marker gene data analysis methods, i.e., ASVs and OTUs, to accurately measure their diversity and applicability to communities. ASVs controlled errors sufficiently, and the resulting OTUs were regarded as the biological counterpart to a species in improved resolution (Callahan et al., 2017 ). Subsequently, 1287 OTUs were annotated at the family level using the NCBI nt database, and those with an abundance greater than 0.1% were classified into 1 of 4 feeding guilds based on the literature, i.e., detritivores, herbivores, microbivores, and predators (Table S2) (Wei et al., 2021 ). 2.5 Data analyses The diversity indices (species richness, Simpson index, and Shannon–Wiener) (Thukral, 2017 ) and community composition (species distribution in each sample) of soil fauna were determined based on the occurrence of each OTU (= species) in individual samples. All statistical analyses were performed using R version 4.3.2 (Team, 2013). The diversity of the soil fauna community was computed using the R packages “picante” and “vegan” (Shannon, 2001 ; Oksanen et al., 2013a ). Due to the non-normal data distribution, we employed the non-parametric Kruskal–Wallis one-way analysis of variance, as implemented in the R package “rstatix” (Shannon, 2001 ; Oksanen et al., 2013a ) to assess the impact of reciprocal litter transplant on soil fauna diversity (Daniel, 1990 ). Furthermore, we performed principal coordinate analysis (PCoA) based on the Bray–Curtis distance using the R packages “vegan” and “ape” (Anderson and Willis, 2003 ; Edwards et al., 2015 ) to identify differences in community composition, dominant classes, and feeding guilds in soil fauna. The “ggplot2” (Wickham, 2011 ) package was used to visualize the PCoA plot. The “adespatial” package (Shen et al., 2020 ) was used to analyze the community and elucidate the factors driving changes in soil fauna community composition through three key processes: species replacement, richness differences, and Jaccard similarity. An UpSet plot was generated using the “UpSetR” package to depict the shared and unique OTUs among treatment groups (Conway et al., 2017 ). Redundancy analysis (RDA) was performed using the “vegan” package (Oksanen et al., 2013b ) and “ggplot2” (Wickham, 2011 ) to explore the correlation between soil fauna and environmental constraints (physicochemical litter properties). 3. Results 3.1 Responses of soil fauna diversity to reciprocal litter transplant In both bamboo and broadleaf forests, the diversity of soil fauna in both the litter and soil layer was not significantly influenced by litter retention, removal, or replacement (exchange) treatments (Fig. S3, P > 0.05). Similarly, in both bamboo and broadleaf forests, soil fauna diversity showed no significant response to litter treatments in the soil layer (Fig. S3, P > 0.05). However, significant differences were observed in the diversity indices of the soil fauna communities between the litter and the soil layer (Fig. S3, P < 0.05). 3.2 Responses of soil fauna community composition to reciprocal litter transplant The community composition of total soil fauna in both the litter and soil layer was significantly affected by reciprocal litter transplant (Table 1 , Fig. 1 a, P < 0.001, Fig. 2 a, P < 0.01). Specifically, reciprocal litter transplant influenced the community compositions of Arachnida, Collembola, and Insecta in the litter layer, but no effect was detected in the soil layer (Table 1 ). Within the litter layer, significant distinctions were observed among all treatments regarding soil fauna communities (Table 2 a, P < 0.001; b, P < 0.001; c, P < 0.05; d, P < 0.01; e, P < 0.001). Table 1 Results of permutational multivariate analysis of variance (PERMANOVA) testing (F-values) the effects of reciprocal litter transplant on the community compositions of total soil fauna, dominant classes, and feeding guilds. Groups Total soil fauna Dominant Classes Feeding guilds Arachnida Collembola Insecta Detritivores Herbivores Microbivores Predators Litter layer 0.0913*** 0.1021*** 0.0981*** 0.0788*** 0.1372** 0.1149** 0.1182** 0.1174*** Soil layer 0.0793** 0.0927 0.1275 0.0745 0.1031 0.1284 0.0713 0.1081** Note: *, P < 0.05; **, P < 0.01; ***, P 0.05), whereas a difference was observed in the broadleaf forest (Table 2 g, P 0.05; j, P 0.05; l, P > 0.05; n, P > 0.05). In contrast, in the broadleaf forest, bamboo litter had a significant impact on the soil fauna community structure (Table 2 g, P 0.05; m, P < 0.01). 3.3 Responses of soil fauna feeding guilds to reciprocal litter transplant Across the four feeding guilds, soil fauna within the soil layer demonstrated lower susceptibility to reciprocal litter transplant compared to those in the litter layer since no effects were observed on herbivores, microbivores, or detritivores in the soil layer (Table 1 , P > 0.05). However, predators were notably affected by reciprocal litter transplant in both the litter and the soil layer (Table 1 , P < 0.01; Fig. 1 h, P < 0.001; Fig. 2 b, P < 0.001). RDA revealed that the feeding guilds of soil fauna within the litter layer had varied correlations with the physicochemical properties of litter (Fig. 3 ). Herbivores exhibited a positive correlation with litter lignin and total nitrogen contents but a negative correlation with the cellulose content. Microbivores showed a positive correlation with the litter cellulose content and the carbon-to-nitrogen ratio. Predators and detritivores demonstrated a positive correlation with the total carbon content. 3.4 Species replacement occurred in soil fauna community composition The ternary plots demonstrated that the total soil fauna communities primarily underwent species replacement (Fig. 4 a and c) with reciprocal litter transplant, but such a distinct replacement process was not detected in the feeding guild communities (Fig. 4 b and d). This result was consistent with that from the UpSet plot (Fig. 5 ), which showed a pattern of high species exclusivity and low species sharing in the overall soil fauna community. Beta diversity decomposition analyses showed that the soil fauna community composition dissimilarities within the litter and the soil layer were primarily driven by species replacement processes, contributing 79.73 and 77.62%, respectively. In contrast, richness difference processes contributed only 20.27 and 22.38%, respectively, on average (Fig. 4 a and c, Table S3). 4. Discussion This study uncovered how bamboo invasion affected soil fauna communities by altering litter quality, thus impacting the ecosystem structure, and provided direct evidence of the close link between soil fauna and aboveground systems. It also showed that despite species turnover in soil fauna, ecological functions, such as decomposition, remained stable, highlighting ecosystem resilience and informing strategies to manage invasive plants without disrupting key ecological processes. We investigated the impact of reciprocal litter transplant on soil fauna diversity and community composition in the context of bamboo invasion, addressing three scientific questions outlined in the experimental design: (1) Litter reciprocal transplant did not significantly affect soil fauna diversity but did alter community composition; (2) changes in soil fauna communities occurred through species replacement without affecting their ecological functions; and (3) changes in soil fauna communities were driven by higher litter quality. 4.1 Litter removal did not affect soil fauna community composition This study demonstrated that soil fauna diversity and community composition in the soil layer did not exhibit significant changes after litter removal compared to litter retention (Table 2 k and n). Similarly, Ashford et al. ( 2013 ) found that litter removal did not alter soil fauna diversity or community biomass. The resilience of soil fauna may be attributed to the communities residing in deeper soil layers being less influenced by external environmental factors (Scherber et al., 2010 ; Yin et al., 2019 ). Long et al. ( 2023 ) found that surface-dwelling collembola were more susceptible to external environmental influences than their soil-dwelling counterparts. In line with this report, our results showed that the soil fauna community structure in the soil layer was less sensitive to reciprocal litter transplant than that in the litter layer (Table 1 ). In other words, local litter did not affect the soil fauna community structure in the soil layer. 4.2 The exchange of litter inputs changed soil fauna community composition This study revealed that reciprocal litter transplant did not affect soil fauna diversity in the context of bamboo invasion (Fig. S3). This finding aligns with previous controlled experiments, which showed no significant differences in soil fauna richness and density across three forest types in a litter decomposition study (Yang and Chen, 2009 ). Furthermore, Ashford et al. ( 2013 ) demonstrated that neither the removal nor the addition of leaf litter significantly affected taxonomic richness or diversity. However, observational investigations have shown that litter inputs from different forest types have significant effects on soil fauna richness and abundance (Cifuentes-Croquevielle et al., 2020 ; Wei et al., 2021 ). These contrasting results may be attributed to the varied conditions between the experiments and the natural forests, highlighting the complexity of ecological interactions in forest ecosystems. The lack of significant effects of reciprocal litter transplant on soil fauna diversity could be explained by ecological functional redundancy. However, soil fauna community composition showed significant alterations after reciprocal litter transplant (Table 1 ). Notably, in the litter layer, significant differences were observed in soil fauna communities between litter retention and replacement treatments (Table 2 a–e), indicating the adaptation of local soil communities to the specific litter type in the decomposition process (Bocock et al., 1960 ; Wardle, 2002 ; Ayres et al., 2009b ; Barantal et al., 2011 ; Austin et al., 2014 ; Zhu et al., 2024 ). The attributes of leaf litter and the forest floor, which act as key food sources and habitats, may affect the composition of soil fauna communities both directly, through providing food, and indirectly, by influencing the soil environment as a habitat (Peng et al., 2022 ). Studies have consistently demonstrated that soil fauna community composition varies across regions dominated by different plant species (Griffiths et al., 1992 ; Grayston et al., 1998 ; Bardgett and Walker, 2004 ; Wei et al., 2021 ). The soil fauna community composition for decomposing the same litter type differed significantly among forest stands (Table 2 d and e). This is likely due to community variations in the native soil fauna for each local stand (Wei et al., 2021 ), with those fauna capable of successful decomposition adopting the specific manipulated litter in their forest stands. Similarly, soil fauna communities within the same forest stand showed variations among litter types (Table 2 b and c), which may be attributed to specific litter types requiring decomposer communities with particular breakdown functions (Hunt et al., 1988 ). Moreover, Li et al. ( 2020 ) found that soil microorganisms specialized in decomposing specific resources at different litter decomposition stages. Such specificity in adaptation allows soil organisms to effectively utilize the litter from particular plant species, which may, in turn, influence organic material cycling and the dynamic balance within soil ecosystem functions. 4.3 Litter quality drives changes in soil fauna communities In this study’s pairwise comparison of three reciprocal litter replacement manipulations, a significant difference in soil-dwelling fauna was only detected when bamboo litter was placed in the broadleaf forest (Table 2 ). Moreover, in the litter layer, the influence of bamboo litter on soil fauna communities was more pronounced than that of broadleaf litter (Table 2 ). This may be related to the high quality of bamboo litter, as it has an elevated N concentration and low lignin/N ratio (Table S1 ) (Phillips et al., 2013 ). Studies have shown that high-quality litter (e.g., high N content and low lignin, phenol, and tannin contents) (Rai et al., 2016 ; Lin et al., 2019 ) ensure greater availability of essential nutrients, such as phosphorus, calcium, and magnesium, in the forest floor, which are critical factors influencing soil fauna communities (Yang et al., 2020 ; Peng et al., 2022 ). Specifically, high-quality litter are harder to decompose (Melillo et al., 1982 ; Vivanco and Austin, 2008 ; Coq et al., 2010 ) and may stimulate native decomposer communities more effectively. In low-quality broadleaf forests, the presence of higher quality bamboo litter significantly altered the soil fauna community composition. In other words, high-quality litter input had a greater impact on the soil fauna community in low-quality litter environments but not vice versa. This finding confirms that leaf litter quality is likely the primary factor behind the effect of tree species on the soil fauna community (Peng et al., 2022 ). The results align with those of previous studies indicating that the decomposer community's capacity to adapt to varying litter qualities may be limited by its adaptation to locally prevalent, low-quality litter (Austin et al., 2014 ; Zhou et al., 2020 ). This phenomenon may be due to the enhanced decomposition rate of bamboo litter, which is facilitated by shifts in soil biota and its higher quality compared to native tree species, thus aiding the invasion process (Luan et al., 2021 ; Liu et al., 2024 ). This could be a successful strategy for bamboo invasion. 4.4 Reciprocal litter transplant altered the soil fauna community through species replacement without changing its ecological function This study found that alterations in soil fauna community composition were primarily driven by the species replacement process, which is consistently observed both in the litter and the soil layer (Fig. 4 a and c). This aligns with the predictions of Filgueiras et al. ( 2021 ), who suggested that ecological groups are predictably replaced as land-use intensifies, which significantly alters habitat availability and quality. Additionally, species replacement can be attributed to soil biological communities associated with specific plant species that specialize in decomposing litter from those plants (Bocock et al., 1960 ; Ayres et al., 2009b ; Strickland et al., 2009 ; Li et al., 2020 ). Consistent with previous research, the high silicon content of bamboo fosters the development of specific decomposer communities during the decomposition process (Schaller et al., 2016 ). Consequently, the litter input changes in our study might lead to corresponding alternations in the soil fauna associated with them. However, the feeding guild communities (trophic levels) did not show a replacement process (Fig. 4 b and d), indicating the stability of the forest soil trophic structure with reciprocal litter transplant. Soil fauna diversity varied greatly between native bamboo forests and broadleaf forests in subtropical regions (Wei et al., 2021 ; Long et al., 2023 ), potentially influencing their capacity for litter decomposition. The ability of soil fauna to decompose litter is not solely determined by their diversity but also by litter quality. When litter decompose in a new environment, litter quality changes alter the associated food web. Although the necessary soil fauna species may be absent, other species with similar feeding habits can fulfill this role, demonstrating ecological functional redundancy, which helps maintain ecosystem stability (Biggs et al., 2020 ). In our study, among treatment groups, the OTUs that were replaced still belonged to the same feeding guild (Table S4). These findings align with those of Sauvadet et al. ( 2017 ), who found that litters with distinct qualities primarily influenced the first decomposer communities without significantly affecting indirect trophic links. In summary, species replacement in soil fauna communities primarily involves substitution with species in the same ecological niche, maintaining ecosystem stability. 5. Conclusion This study highlights how plant invasion influences the soil fauna community through alterations in litter quality. Specifically, reciprocal litter transplant during bamboo invasions reshaped soil fauna communities through species replacement. Notably, introducing high-quality litter into low-quality litter stands triggered substantial changes in the soil fauna community. Despite these shifts, the trophic structure of the soil fauna remained stable, reflecting the resilience and functional stability of local soil ecosystems. This further confirms the strong relationship between soil fauna and aboveground systems. These insights provide a potential strategy for managing plant invasions by modifying litter inputs to influence soil fauna communities. Such an approach can suppress invasive plants and preserve essential ecological functions, such as organic matter decomposition and nutrient cycling. The complex relationship between soil fauna and litter quality plays a pivotal role in maintaining ecosystem health and offers promising avenues for ecosystem management. Declarations CRediT authorship contribution statement Kui Long : Data curation, Formal analysis, Methodology, Software, Validation, Visualization, Writing – original draft, Writing – review & editing. Zhenyu Zhou : Data curation, Formal analysis, Investigation, Validation, Visualization, Writing – original draft. Gan Yu : Data curation, Investigation, Formal analysis. Yakun Zhang : Writing – review & editing. Yang Wang : Investigation, Formal analysis. Qingyun Wang : Formal analysis, Project administration, Software. Yongchun Li : Conceptualization, Funding acquisition, Project administration, Resources, Writing – review & editing. Junhao Huang : Conceptualization, Funding acquisition, Methodology, Project administration, Resources, Supervision, Writing – review & editing. All authors validated the findings and approved the final version for publication. Data availability Illumina sequence data supporting the results of this study will be submitted to NCBI once paper is accepted by the journal. Declaration of interests The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper. Acknowledgements The authors are very grateful to Professor Feng Zhang (Nanjing Agricultural University), Dr. Xiaolong Lin (Shanghai Ocean University), and Professor Jianjun Guo (Guizhou University) for their great support for soil fauna identifications. The authors also thank Dr. Bismillah Shah, Ms. Yi Zhu, Mr. Ao Li, and Mr. Yisa Shao (Zhejiang A&F University, Hangzhou, China) for their assistance with fieldwork. This study was supported by the National Natural Science Foundation of China (NSFC, Grant No. 32071742). Declaration of generative AI and AI-assisted technologies in the writing process During the preparation of this work the authors used ChatGPT in order to improve the language and readability of the manuscript. After using this tool, the authors reviewed and edited the content as needed and take full responsibility for the content of the publication. References Anderson, M.J., Willis, T.J., 2003. Canonical analysis of principal coordinates: a useful method of constrained ordination for ecology. Ecology 84, 511-525. https://doi.org/10.1890/0012-9658(2003)084[0511:CAOPCA]2.0.CO;2 Arribas, P., Andújar, C., Hopkins, K., Shepherd, M., Vogler, A.P., 2016. Metabarcoding and mitochondrial metagenomics of endogean arthropods to unveil the mesofauna of the soil. Methods Ecol. Evol. 7, 1071-1081.https://doi.org/10.1111/2041-210X.12557 Ashford, O., Foster, W., Turner, B.L., Sayer, E., Sutcliffe, L., Tanner, E., 2013. Litter manipulation and the soil arthropod community in a lowland tropical rainforest. Soil Biol. Biochem. 62, 5-12. https://doi.org/10.1016/j.soilbio.2013.03.001 Austin, A.T., Vivanco, L., González‐Arzac, A., Pérez, L.I., 2014. There's no place like home? An exploration of the mechanisms behind plant litter–decomposer affinity in terrestrial ecosystems. Wiley Online Library.https://doi.org/10.1111/nph.12959 Ayres, E., Steltzer, H., Berg, S., Wall, D.H., 2009a. Soil biota accelerate decomposition in high‐elevation forests by specializing in the breakdown of litter produced by the plant species above them. J. Ecol. 97, 901-912. https://doi.org/10.1111/j.1365-2745.2009.01539.x Ayres, E., Steltzer, H., Berg, S., Wallenstein, M.D., Simmons, B.L., Wall, D.H., 2009b. Tree species traits influence soil physical, chemical, and biological properties in high elevation forests. PloS one 4, e5964. https://doi.org/10.1371/journal.pone.0005964 Bacher, S., Blackburn, T.M., Essl, F., Genovesi, P., Heikkilä, J., Jeschke, J.M., Jones, G., Keller, R., Kenis, M., Kueffer, C., 2018. Socio‐economic impact classification of alien taxa (SEICAT). Methods Ecol. Evol. 9, 159-168. https://doi.org/10.1111/2041-210X.12844 Balasubramanian, A., Sivakumar, B., CN, H.P., Swathiga, G., Vasanth, V., Thirumoorthy, P., Bora, N.R., SJ, M.P., 2023. Influence of Soil Invertebrates on Soil Decomposition. J. Surv. Fish. Sci. 10, 618-629. https://doi.org/10.53555/sfs.v10i3.1815 Barantal, S., Roy, J., Fromin, N., Schimann, H., Hättenschwiler, S., 2011. Long-term presence of tree species but not chemical diversity affect litter mixture effects on decomposition in a neotropical rainforest. Oecologia 167, 241-252. https://doi.org/10.1007/s00442-011-1966-4 Bardgett, R.D., Chan, K.F., 1999. Experimental evidence that soil fauna enhance nutrient mineralization and plant nutrient uptake in montane grassland ecosystems. Soil Biol. Biochem. 31, 1007-1014. https://doi.org/10.1016/S0038-0717(99)00014-0 Bardgett, R.D., Walker, L.R., 2004. Impact of coloniser plant species on the development of decomposer microbial communities following deglaciation. Soil Biol. Biochem. 36, 555-559. https://doi.org/10.1016/j.soilbio.2003.11.002 Biggs, C.R., Yeager, L.A., Bolser, D.G., Bonsell, C., Dichiera, A.M., Hou, Z., Keyser, S.R., Khursigara, A.J., Lu, K., Muth, A.F., 2020. Does functional redundancy affect ecological stability and resilience? A review and meta-analysis. Ecosphere 11, e03184. https://doi.org/10.1002/ecs2.3184 Bocock, K., Gilbert, O., Capstick, C., Twinn, D., Waid, J., Woodman, M., 1960. Changes in leaf litter when placed on the surface of soils with contrasting humus types: I. Losses in dry weight of oak and ash leaf litter. J. Soil Sci. 11, 1-9. https://doi.org/10.1111/j.1365-2389.1960.tb02196.x Bolger, A.M., Lohse, M., Usadel, B., 2014. Trimmomatic: a flexible trimmer for Illumina sequence data. Bioinformatics 30, 2114-2120. 10.1093/bioinformatics/btu170 Brown, G.G., 1995. How do earthworms affect microfloral and faunal community diversity?, The Significance and Regulation of Soil Biodiversity: Proceedings of the International Symposium on Soil Biodiversity, held at Michigan State University, East Lansing, May 3–6, 1993. Springer, pp. 247-269. https://doi.org/10.1007/BF02183068 Brussaard, L., 1998. Soil fauna, guilds, functional groups and ecosystem processes. Appl. Soil Ecol. 9, 123-135. https://doi.org/10.1016/S0929-1393(98)00066-3 Callahan, B.J., McMurdie, P.J., Holmes, S.P., 2017. Exact sequence variants should replace operational taxonomic units in marker-gene data analysis. ISME J. 11, 2639-2643. https://doi.org/10.1038/ismej.2017.119 Cavallet, B.V., Silva, E.R., Baretta, C.R.D.M., Rezende, R.d.S., 2022. Effect of agriculture land use on standard cellulosic substrates breakdown and invertebrates’ community. Community Ecol. 23, 277-288. https://doi.org/10.1007/s42974-022-00103-9 Chapin, F.S., Matson, P.A., Mooney, H.A., Vitousek, P.M., 2002. Principles of terrestrial ecosystem ecology. https://doi.org/10.1007/b97397 Chen, Z., Li, Y., Chang, S.X., Xu, Q., Li, Y., Ma, Z., Qin, H., Cai, Y., 2021. Linking enhanced soil nitrogen mineralization to increased fungal decomposition capacity with Moso bamboo invasion of broadleaf forests. Sci. Total Environ. 771, 144779. https://doi.org/10.1016/j.scitotenv.2020.144779 Cifuentes-Croquevielle, C., Stanton, D.E., Armesto, J.J., 2020. Soil invertebrate diversity loss and functional changes in temperate forest soils replaced by exotic pine plantations. Sci. Rep. 10, 7762. https://doi.org/10.1038/s41598-020-64453-y Coleman, D.C., Callaham, M.A., Crossley Jr, D., 2017. Fundamentals of soil ecology. Academic press. https://doi.org/10.1016/C2015-0-04083-7 Coleman, D.C., Geisen, S., Wall, D., 2024. Soil fauna: Occurrence, biodiversity, and roles in ecosystem function, Soil microbiology, ecology and biochemistry. Elsevier, pp. 131-159. https://doi.org/10.1016/B978-0-12-822941-5.00005-3 Conway, J.R., Lex, A., Gehlenborg, N., 2017. UpSetR: an R package for the visualization of intersecting sets and their properties. Bioinformatics 33, 2938-2940. https://doi.org/10.1093/bioinformatics/btx364 Coq, S., Souquet, J.-M., Meudec, E., Cheynier, V., Hättenschwiler, S., 2010. Interspecific variation in leaf litter tannins drives decomposition in a tropical rain forest of French Guiana. Ecology 91, 2080-2091. https://doi.org/10.1890/09-1076.1 Creedy, T.J., Ng, W.S., Vogler, A.P., 2019. Toward accurate species‐level metabarcoding of arthropod communities from the tropical forest canopy. Ecol. Evol. 9, 3105-3116. https://doi.org/10.1002/ece3.4839 Daniel, W.W., 1990. Kruskal–Wallis one-way analysis of variance by ranks. Applied nonparametric statistics, 226-234. David, P., Thebault, E., Anneville, O., Duyck, P.-F., Chapuis, E., Loeuille, N., 2017. Impacts of invasive species on food webs: a review of empirical data. Adv. Ecol. Res. 56, 1-60. https://doi.org/10.1016/bs.aecr.2016.10.001 Edwards, J., Johnson, C., Santos-Medellín, C., Lurie, E., Podishetty, N.K., Bhatnagar, S., Eisen, J.A., Sundaresan, V., 2015. Structure, variation, and assembly of the root-associated microbiomes of rice. Proc. Natl. Acad. Sci. U.S.A. 112, E911-E920. https://doi.org/10.1073/pnas.1414592112 Ehrenfeld, J.G., 2010. Ecosystem consequences of biological invasions. Annu. Rev. Ecol. Evol. Syst. 41, 59-80. https://doi.org/10.1146/annurev-ecolsys-102209-144650 Faber, J.H., Verhoef, H.A., 1991. Functional differences between closely-related soil arthropods with respect to decomposition processes in the presence or absence of pine tree roots. Soil Biol. Biochem. 23, 15-23. https://doi.org/10.1016/0038-0717(91)90157-F Filgueiras, B.K., Peres, C.A., Melo, F.P., Leal, I.R., Tabarelli, M., 2021. Winner–loser species replacements in human-modified landscapes. Trends Ecol. Evol. 36, 545-555. https://doi.org/10.1016/j.tree.2021.02.006 Fukushima, K., Usui, N., Ogawa, R., Tokuchi, N., 2015. Impacts of moso bamboo (Phyllostachys pubescens) invasion on dry matter and carbon and nitrogen stocks in a broad‐leaved secondary forest located in K yoto, western J apan. Plant Species Biol. 30, 81-95. https://doi.org/10.1111/1442-1984.12066 Gallardo, B., Bacher, S., Barbosa, A.M., Gallien, L., González-Moreno, P., Martínez-Bolea, V., Sorte, C., Vimercati, G., Vilà, M., 2024. Risks posed by invasive species to the provision of ecosystem services in Europe. Nat. Commun. 15, 2631. https://doi.org/10.1038/s41467-024-46818-3 Grayston, S.J., Wang, S., Campbell, C.D., Edwards, A.C., 1998. Selective influence of plant species on microbial diversity in the rhizosphere. Soil Biol. Biochem. 30, 369-378. https://doi.org/10.1016/S0038-0717(97)00124-7 Griffiths, B., Welschen, R.v., Van Arendonk, J., Lambers, H., 1992. The effect of nitrate-nitrogen supply on bacteria and bacterial-feeding fauna in the rhizosphere of different grass species. Oecologia 91, 253-259. https://doi.org/10.1007/BF00317793 Heděnec, P., Jiménez, J.J., Moradi, J., Domene, X., Hackenberger, D., Barot, S., Frossard, A., Oktaba, L., Filser, J., Kindlmann, P., 2022. Global distribution of soil fauna functional groups and their estimated litter consumption across biomes. Sci. Rep. 12, 17362. https://doi.org/10.1038/s41598-022-21563-z Hunt, H., Ingham, E., Coleman, D., Elliott, E., Reid, C., 1988. Nitrogen limitation of production and decomposition in prairie, mountain meadow, and pine forest. Ecology 69, 1009-1016. https://doi.org/10.2307/1941256 Jones, T., Thompson, L., Lawton, J., Bezemer, T., Bardgett, R., Blackburn, T., Bruce, K., Cannon, P., Hall, G., Hartley, S., 1998. Impacts of rising atmospheric carbon dioxide on model terrestrial ecosystems. Science 280, 441-443. https://doi.org/10.1126/science.280.5362.441 Kappes, H., Lay, R., Topp, W., 2007. Changes in different trophic levels of litter-dwelling macrofauna associated with giant knotweed invasion. Ecosystems 10, 734-744. https://doi.org/10.1007/s10021-007-9052-9 Kassambara, A., 2019. Comparing groups: Numerical variables. Datanovia.[Google Scholar]. Khattak, W.A., Sun, J., Hameed, R., Zaman, F., Abbas, A., Khan, K.A., Elboughdiri, N., Akbar, R., He, F., Ullah, M.W., 2024. Unveiling the resistance of native weed communities: insights for managing invasive weed species in disturbed environments. Biol. Rev. 99, 753-777. https://doi.org/10.1111/brv.13043 Li, C., Bo, H., Song, B., Chen, X., Cao, Q., Yang, R., Ji, S., Wang, L., Liu, J., 2022. Reshaping of the soil microbiome by the expansion of invasive plants: shifts in structure, diversity, co-occurrence, niche breadth, and assembly processes. Plant Soil 477, 629-646. https://doi.org/10.1007/s11104-022-05445-x Li, D., Wei, J., Wu, J., Zhong, Y., Chen, Z., He, J., Zhang, S., Yu, L., 2024. The expansion of moso bamboo (Phyllostachys edulis) forests into diverse types of forests in China from 2010 to 2020. Forests 15, 1418. https://doi.org/10.3390/f15081418 Li, Y., Ma, L., Wang, J., Zhang, J., 2021. Soil faunal community composition alters nitrogen distribution in different land use types in the Loess Plateau, China. Appl. Soil Ecol. 163, 103910. https://doi.org/10.1016/j.apsoil.2021.103910 Li, Y., Veen, G.C., Hol, W.G., Vandenbrande, S., Hannula, S.E., ten Hooven, F.C., Li, Q., Liang, W., Bezemer, T.M., 2020. ‘Home’and ‘away’litter decomposition depends on the size fractions of the soil biotic community. Soil Biol. Biochem. 144, 107783. https://doi.org/10.1016/j.soilbio.2020.107783 Liao, C., Peng, R., Luo, Y., Zhou, X., Wu, X., Fang, C., Chen, J., Li, B., 2008. Altered ecosystem carbon and nitrogen cycles by plant invasion: a meta‐analysis. New Phytol. 177, 706-714. https://doi.org/10.1111/j.1469-8137.2007.02290.x Lieurance, D., Cooper, A., Young, A.L., Gordon, D.R., Flory, S.L., 2018. Running bamboo species pose a greater invasion risk than clumping bamboo species in the continental United States. J. Nat. Conserv. 43, 39-45. https://doi.org/10.1016/j.jnc.2018.02.012 Lin, D., Pang, M., Fanin, N., Wang, H., Qian, S., Zhao, L., Yang, Y., Mi, X., Ma, K., 2019. Fungi participate in driving home-field advantage of litter decomposition in a subtropical forest. Plant Soil 434, 467-480. https://doi.org/10.1007/s11104-018-3865-5 Liu, J., Compson, Z.G., Gui, X., Yang, Q., Song, Q., Huang, D., Ren, Z., Luan, F., 2024. Weak responses of soil microorganisms to leaf litter inputs after native Phyllostachys edulis invasion into adjacent native forests. Plant Soil 494, 685-699. https://doi.org/10.1007/s11104-023-06311-0 Liu, X., Siemann, E., Cui, C., Liu, Y., Guo, X., Zhang, L., 2019. Moso bamboo (Phyllostachys edulis) invasion effects on litter, soil and microbial PLFA characteristics depend on sites and invaded forests. Plant Soil 438, 85-99. https://doi.org/10.1007/s11104-019-04010-3 Long, K., Yin, R., Kardol, P., Wei, Q., Li, Y., Huang, J., 2023. Bamboo invasion alters Collembola community composition varying with life‐forms. Pest Manag. Sci. 79, 2517-2526. https://doi.org/10.1002/ps.7434 Luan, J., Liu, S., Li, S., Whalen, J.K., Wang, Y., Wang, J., Liu, Y., Dong, W., Chang, S.X., 2021. Functional diversity of decomposers modulates litter decomposition affected by plant invasion along a climate gradient. J. Ecol. 109, 1236-1249. https://doi.org/10.1111/1365-2745.13548 Ma, A., Liu, H., Song, C., Tian, E., Wang, X., 2023. Home‐field advantage in litter decomposition: A critical review from a microbial perspective. J. Basic Microbiol. 63, 709-721. https://doi.org/10.1002/jobm.202200657 Macfadyen, A., 1953. Notes on methods for the extraction of small soil arthropods. J. Anim. Ecol., 65-77. https://doi.org/10.1016/S0031-4056(23)00341-4 Magoč, T., Salzberg, S.L., 2011. FLASH: fast length adjustment of short reads to improve genome assemblies. Bioinformatics 27, 2957-2963. https://doi.org/10.1093/bioinformatics/btr507 Melillo, J.M., Aber, J.D., Muratore, J.F., 1982. Nitrogen and lignin control of hardwood leaf litter decomposition dynamics. Ecology 63, 621-626. https://doi.org/10.2307/1936780 Milcu, A., Manning, P., 2011. All size classes of soil fauna and litter quality control the acceleration of litter decay in its home environment. Oikos 120, 1366-1370. https://doi.org/10.1111/j.1600-0706.2010.19418.x Morris, S.J., Blackwood, C.B., 2024. The ecology of soil biota and their function, Soil microbiology, ecology and biochemistry. Elsevier, pp. 275-302. https://doi.org/10.1016/B978-0-12-822941-5.00010-7 Negrete-Yankelevich, S., Fragoso, C., Newton, A.C., Russell, G., Heal, O.W., 2008. Species-specific characteristics of trees can determine the litter macroinvertebrate community and decomposition process below their canopies. Plant Soil 307, 83-97. https://doi.org/10.1007/s11104-008-9585-5 Oksanen, J., Blanchet, F., Kindt, R., Legendre, P., Minchin, P., O’hara, R., Simpson, G., Solymos, P., Stevens, M., Wagner, H., 2013a. Package ‘vegan’. Community Ecol. package, version 2. Oksanen, J., Blanchet, F.G., Kindt, R., Legendre, P., Minchin, P.R., O’hara, R., Simpson, G.L., Solymos, P., Stevens, M.H.H., Wagner, H., 2013b. Package ‘vegan’. Community Ecol. package, version 2, 1-295. Olivia, C., Dumitru, M.T., 2023. THE FUNCTIONAL GROUPS OF INVERTEBRATES ACCORDING TO THEIR TROPHIC PREFERENCE. Oltenia, Stud. Comun. Stiintele Nat. 39. Peng, Y., Holmstrup, M., Schmidt, I.K., De Schrijver, A., Schelfhout, S., Heděnec, P., Zheng, H., Bachega, L.R., Yue, K., Vesterdal, L., 2022. Litter quality, mycorrhizal association, and soil properties regulate effects of tree species on the soil fauna community. Geoderma 407, 115570. https://doi.org/10.1016/j.geoderma.2021.115570 Peng, Y., Schmidt, I.K., Zheng, H., Heděnec, P., Bachega, L.R., Yue, K., Wu, F., Vesterdal, L., 2020. Tree species effects on topsoil carbon stock and concentration are mediated by tree species type, mycorrhizal association, and N-fixing ability at the global scale. Forest Ecol. Manag. 478, 118510. https://doi.org/10.1016/j.foreco.2020.118510 Peng, Z., Lu, Y., Li, L., Zhao, Q., Feng, Q., Gao, Z., Lu, H., Hu, T., Yao, N., Liu, K., 2013. The draft genome of the fast-growing non-timber forest species moso bamboo (Phyllostachys heterocycla). Nat. Genet. 45, 456-461. https://doi.org/10.1038/ng.2569 Pérez, G., Vila, M., Gallardo, B., 2022. Potential impact of four invasive alien plants on the provision of ecosystem services in Europe under present and future climatic scenarios. Ecosyst. Serv. 56, 101459. https://doi.org/10.1016/j.ecoser.2022.101459 Petersen, H., 1982. Basis for evaluation of the density and biomass data. Oikos 39, 295-305. Phillips, R.P., Brzostek, E., Midgley, M.G., 2013. The mycorrhizal‐associated nutrient economy: a new framework for predicting carbon–nutrient couplings in temperate forests. New Phytol. 199, 41-51. https://doi.org/10.1111/nph.12221 Premakumari, P.D., Sarayu, M.G., Das, G.G.M., Babu, K.V.D., Krishnan, R., Lawarence, B., Siraj, S.S., Pillai, A.R., Syamala, S.K.S., Murugan, K., 2022. Invasive exotic plant species and their influence on the environment, ecosystem services, economy and health: a search. J. Adv. Sci. Res. 13, 64-74. https://doi.org/10.55218/JASR.202213106 Raheem, A., Yohanna, P., Li, G., Noh, N.J., Iqbal, B., Tang, J., Du, D., Alahmadi, T.A., Ansari, M.J., Zhan, A., 2024. Unraveling the ecological threads: How invasive alien plants influence soil carbon dynamics. J. Environ. Manage. 356, 120556. https://doi.org/10.1016/j.jenvman.2024.120556 Rai, A., Singh, A.K., Ghosal, N., Singh, N., 2016. Understanding the effectiveness of litter from tropical dry forests for the restoration of degraded lands. Ecol. Eng. 93, 76-81. https://doi.org/10.1016/j.ecoleng.2016.05.014 Rai, P.K., Singh, J., 2020. Invasive alien plant species: Their impact on environment, ecosystem services and human health. Ecol. Indic. 111, 106020. 10.1016/j.ecolind.2019.106020 Reddy, H.R., Khan, M., Pasarkar, A., 2024. Soil Invertebrates As Sentinels Of Soil Health: A Zoological Approach To Soil Quality Assessment. Journal of Advanced Zoology 45. https://doi.org/10.53555/jaz.v45i1.3007 Sauvadet, M., Chauvat, M., Brunet, N., Bertrand, I., 2017. Can changes in litter quality drive soil fauna structure and functions? Soil Biol. Biochem. 107, 94-103. https://doi.org/10.1016/j.soilbio.2016.12.018 Schaller, J., Böttger, R., Dudel, G., Ruess, L., 2016. Reed litter Si content affects microbial community structure and the lipid composition of an invertebrate shredder during aquatic decomposition. Limnologica 57, 14-22. https://doi.org/10.1016/j.limno.2015.12.002 Scherber, C., Eisenhauer, N., Weisser, W.W., Schmid, B., Voigt, W., Fischer, M., Schulze, E.-D., Roscher, C., Weigelt, A., Allan, E., 2010. Bottom-up effects of plant diversity on multitrophic interactions in a biodiversity experiment. Nature 468, 553-556. https://doi.org/10.1038/nature09492 Seastedt, T., Crossley Jr, D., 1983. Nutrients in forest litter treated with naphthalene and simulated throughfall: a field microcosm study. Soil Biol. Biochem. 15, 159-165. https://doi.org/10.1016/0038-0717(83)90097-4 Setälä, H., Marshall, V.G., Trofymow, J., 1996. Influence of body size of soil fauna on litter decomposition and 15N uptake by poplar in a pot trial. Soil Biol. Biochem. 28, 1661-1675. https://doi.org/10.1016/S0038-0717(96)00252-0 https://doi.org/10.1016/S0038-0717(96)00252-0 Shannon, C.E., 2001. A mathematical theory of communication. ACM SIGMOBILE mobile computing and communications review 5, 3-55. https://doi.org/10.1145/584091.584093 Shen, C., Gunina, A., Luo, Y., Wang, J., He, J.Z., Kuzyakov, Y., Hemp, A., Classen, A.T., Ge, Y., 2020. Contrasting patterns and drivers of soil bacterial and fungal diversity across a mountain gradient. Environ. Microbiol. 22, 3287-3301. https://doi.org/10.1111/1462-2920.15090 Shokralla, S., Porter, T.M., Gibson, J.F., Dobosz, R., Janzen, D.H., Hallwachs, W., Golding, G.B., Hajibabaei, M., 2015. Massively parallel multiplex DNA sequencing for specimen identification using an Illumina MiSeq platform. Sci. Rep. 5, 1-7. https://doi.org/10.1038/srep09687 Strickland, M.S., Osburn, E., Lauber, C., Fierer, N., Bradford, M.A., 2009. Litter quality is in the eye of the beholder: initial decomposition rates as a function of inoculum characteristics. Funct. Ecol. 23, 627-636. https://doi.org/10.1111/j.1365-2435.2008.01515.x Sun, H., Hu, W., Dai, Y., Ai, L., Wu, M., Hu, J., Zuo, Z., Li, M., Yang, H., Ma, J., 2023. Moso bamboo (Phyllostachys edulis (Carrière) J. Houzeau) invasion affects soil microbial communities in adjacent planted forests in the Lijiang River basin, China. Front. Microbiol. 14, 1111498. https://doi.org/10.3389/fmicb.2023.1111498 Team, R.C., 2013. R: A language and environment for statistical computing. Teng, Q., Lu, X., Zhang, Q., Cai, L., Sardar, M.F., Li, Y., Abbas, T., Li, Y., Chang, S.X., Li, Y., 2023. Litterfall quality modulates soil ammonium and nitrate supply through altering microbial function in bamboo encroachment of broadleaf forests. Geoderma 437, 116592. https://doi.org/10.1016/j.geoderma.2023.116592 Thukral, A.K., 2017. A review on measurement of Alpha diversity in biology. Agric. Res. J. 54. https://doi.org/10.5958/2395-146X.2017.00001.1 Verhoef, H., Brussaard, L., 1990. Decomposition and nitrogen mineralization in natural and agroecosystems: the contribution of soil animals. Biogeochemistry 11, 175-211. https://doi.org/10.1007/BF00004496 Vilà, M., Espinar, J.L., Hejda, M., Hulme, P.E., Jarošík, V., Maron, J.L., Pergl, J., Schaffner, U., Sun, Y., Pyšek, P., 2011. Ecological impacts of invasive alien plants: a meta‐analysis of their effects on species, communities and ecosystems. Ecol. Lett. 14, 702-708. https://doi.org/10.1111/j.1461-0248.2011.01628.x Vivanco, L., Austin, A.T., 2008. Tree species identity alters forest litter decomposition through long‐term Plant Soil interactions in Patagonia, Argentina. J. Ecol. 96, 727-736. https://doi.org/10.1111/j.1365-2745.2008.01393.x Wang, J., Wang, Y., He, N., Ye, Z., Chen, C., Zang, R., Feng, Y., Lu, Q., Li, J., 2020. Plant functional traits regulate soil bacterial diversity across temperate deserts. Sci. Total Environ. 715, 136976. https://doi.org/10.1016/j.scitotenv.2020.136976 Wardle, D., 2002. Communities and ecosystems: linking the aboveground and belowground components. https://doi.org/10.1515/9781400847297 Wei, Q., Rui, Y., Huang, J., Vogler, A.P., Li, Y., Miao, X., Kardolg, P., 2021. The diversity of soil mesofauna decline after bamboo invasion in subtropical China. Sci. Total Environ., 147982. https://doi.org/10.1016/j.scitotenv.2021.147982 Weidlich, E.W., Flórido, F.G., Sorrini, T.B., Brancalion, P.H., 2020. Controlling invasive plant species in ecological restoration: A global review. J. Appl. Ecol. 57, 1806-1817. https://doi.org/10.1111/1365-2664.13656 Wickham, H., 2011. ggplot2. Wiley interdisciplinary reviews: computational statistics 3, 180-185. http://dx.doi.org/10.1002/wics.1637 Wolfe, B.E., Klironomos, J.N., 2005. Breaking new ground: soil communities and exotic plant invasion. BioScience 55, 477-487. https://doi.org/10.1641/0006-3568(2005)055[0477:BNGSCA]2.0.CO;2 WRB, 2006. A framework for international classification, correlation and communication. World Soil Resour. Rep. 103. Xiao, H., Liu, J., Li, F., 2023. Both alpha and beta diversity of nematode declines in response to moso bamboo expansion in south China. Appl. Soil Ecol. 183, 104761. https://doi.org/10.1016/j.apsoil.2022.104761 Xu, Q.-F., Liang, C.-F., Chen, J.-H., Li, Y.-C., Qin, H., Fuhrmann, J.J., 2020. Rapid bamboo invasion (expansion) and its effects on biodiversity and soil processes+. Glob. Ecol. Conserv. 21, e00787. https://doi.org/10.1016/j.gecco.2019.e00787 Yang, K., Zhu, J., Zhang, W., Zhang, Q., Lu, D., Zhang, Y., Zheng, X., Xu, S., Wang, G.G., 2022. Litter decomposition and nutrient release from monospecific and mixed litters: Comparisons of litter quality, fauna and decomposition site effects. J. Ecol. 110, 1673-1686. https://doi.org/10.1111/1365-2745.13902 Yang, X., Chen, J., 2009. Plant litter quality influences the contribution of soil fauna to litter decomposition in humid tropical forests, southwestern China. Soil Biol. Biochem. 41, 910-918. https://doi.org/10.1016/j.soilbio.2008.12.028 Yang, Y., Wu, Q., Yang, W., Wu, F., Zhang, L., Xu, Z., Liu, Y., Tan, B., Li, H., Zhou, W., 2020. Temperature and soil nutrients drive the spatial distributions of soil macroinvertebrates on the eastern Tibetan Plateau. Ecosphere 11, e03075. https://doi.org/10.1002/ecs2.3075 Yin, R., Gruss, I., Eisenhauer, N., Kardol, P., Thakur, M.P., Schmidt, A., Xu, Z., Siebert, J., Zhang, C., Wu, G.-L., 2019. Land use modulates the effects of climate change on density but not community composition of Collembola. Soil Biol. Biochem. 138, 107598. https://doi.org/10.1016/j.soilbio.2019.107598 Yin, X., Song, B., Dong, W., Xin, W., Wang, Y., 2010. A review on the eco-geography of soil fauna in China. J. Geogr. Sci. 20, 333-346. https://doi.org/10.1007/s11442-010-0333-4 Yu, D.W., Ji, Y., Emerson, B.C., Wang, X., Ye, C., Yang, C., Ding, Z., 2012. Biodiversity soup: metabarcoding of arthropods for rapid biodiversity assessment and biomonitoring. Methods Ecol. Evol. 3, 613-623. https://doi.org/10.1111/j.2041-210X.2012.00198.x Zhang, L., 2023a. Bamboo Expansion and Soil Microbial Communities, Bamboo Expansion: Processes, Impacts, and Management. Springer, pp. 197-208. Zhang, L., 2023b. Bamboo Expansion Into Adjacent Ecosystems, Bamboo Expansion: Processes, Impacts, and Management. Springer, pp. 19-37. Zhao, C., Li, Y., Zhang, C., Miao, Y., Liu, M., Zhuang, W., Shao, Y., Zhang, W., Fu, S., 2021. Considerable impacts of litter inputs on soil nematode community composition in a young Acacia crassicapa plantation. Soil Ecol. Lett. 3, 145-155. https://doi.org/10.1007/s42832-021-0085-3 Zhong, Y., Yan, W., Wang, R., Wang, W., Shangguan, Z., 2018. Decreased occurrence of carbon cycle functions in microbial communities along with long-term secondary succession. Soil Biol. Biochem. 123, 207-217. https://doi.org/10.1016/j.soilbio.2018.05.017 Zhou, S., Butenschoen, O., Barantal, S., Handa, I.T., Makkonen, M., Vos, V., Aerts, R., Berg, M.P., McKie, B., Van Ruijven, J., 2020. Decomposition of leaf litter mixtures across biomes: The role of litter identity, diversity and soil fauna. J. Ecol. 108, 2283-2297. https://doi.org/10.1111/1365-2745.13452 Zhu, M., Fanin, N., Wang, Q., Xu, Z., Liang, S., Ye, J., Lin, F., Yuan, Z., Mao, Z., Wang, X., 2024. High functional breadth of microbial communities decreases home-field advantage of litter decomposition. Soil Biol. Biochem. 188, 109232. https://doi.org/10.1016/j.soilbio.2023.109232 Table Table 2 is available in the Supplementary Files section. Supplementary Files Supplementalmaterials20250509.docx Table2.docx Cite Share Download PDF Status: Published Journal Publication published 03 Sep, 2025 Read the published version in Plant and Soil → Version 1 posted Editorial decision: Major revisions 13 Jul, 2025 Reviewers agreed at journal 05 Jun, 2025 Reviewers invited by journal 02 Jun, 2025 Editor invited by journal 09 May, 2025 Editor assigned by journal 09 May, 2025 First submitted to journal 08 May, 2025 You are reading this latest preprint version Research Square lets you share your work early, gain feedback from the community, and start making changes to your manuscript prior to peer review in a journal. As a division of Research Square Company, we’re committed to making research communication faster, fairer, and more useful. We do this by developing innovative software and high quality services for the global research community. Our growing team is made up of researchers and industry professionals working together to solve the most critical problems facing scientific publishing. Also discoverable on Platform About Our Team In Review Editorial Policies Advisory Board Help Center Resources Author Services Accessibility API Access RSS feed Manage Cookie Preferences © Research Square 2026 | ISSN 2693-5015 (online) Privacy Policy Terms of Service Do Not Sell My Personal Information {"props":{"pageProps":{"initialData":{"identity":"rs-6625409","acceptedTermsAndConditions":true,"allowDirectSubmit":false,"archivedVersions":[],"articleType":"Research Article","associatedPublications":[],"authors":[{"id":465498657,"identity":"bde6060b-f45f-4e11-92b7-e287f5c4afec","order_by":0,"name":"Kui Long","email":"","orcid":"","institution":"","correspondingAuthor":false,"prefix":"","firstName":"Kui","middleName":"","lastName":"Long","suffix":""},{"id":465498658,"identity":"fb0ddc65-14ea-493e-97d9-41d6483a6e47","order_by":1,"name":"Zhenyu Zhou","email":"","orcid":"","institution":"","correspondingAuthor":false,"prefix":"","firstName":"Zhenyu","middleName":"","lastName":"Zhou","suffix":""},{"id":465498659,"identity":"a7142193-5edd-4d22-9151-f6faa2365187","order_by":2,"name":"Gan Yu","email":"","orcid":"","institution":"","correspondingAuthor":false,"prefix":"","firstName":"Gan","middleName":"","lastName":"Yu","suffix":""},{"id":465498660,"identity":"70bb73b5-968b-4c76-b740-84716baccc2b","order_by":3,"name":"Yakun Zhang","email":"","orcid":"","institution":"","correspondingAuthor":false,"prefix":"","firstName":"Yakun","middleName":"","lastName":"Zhang","suffix":""},{"id":465498661,"identity":"1db46e94-74a0-4267-ba42-525f6b0afc46","order_by":4,"name":"Yang Wang","email":"","orcid":"","institution":"","correspondingAuthor":false,"prefix":"","firstName":"Yang","middleName":"","lastName":"Wang","suffix":""},{"id":465498662,"identity":"919e51b4-3587-4a8d-b78d-6a19dae127e5","order_by":5,"name":"Qingyun Wang","email":"","orcid":"","institution":"","correspondingAuthor":false,"prefix":"","firstName":"Qingyun","middleName":"","lastName":"Wang","suffix":""},{"id":465498663,"identity":"29988bb5-df44-4aa9-b9f2-789deadfe101","order_by":6,"name":"Yongchun Li","email":"","orcid":"","institution":"","correspondingAuthor":false,"prefix":"","firstName":"Yongchun","middleName":"","lastName":"Li","suffix":""},{"id":465498664,"identity":"b0afc85e-6ef7-4b60-8da5-d1c3a113d8f7","order_by":7,"name":"Junhao Huang","email":"data:image/png;base64,iVBORw0KGgoAAAANSUhEUgAAAZAAAAAyAQMAAABI0h/eAAAABlBMVEX///8AAABVwtN+AAAACXBIWXMAAA7EAAAOxAGVKw4bAAAA3UlEQVRIiWNgGAWjYBACAzDJc0COgYHxAZDFTLwWY6BqA1K0MBxIbCBai7lE8rOHX2TupM9vP8wmwVBhndjAfvYAXi2WM9LMjWV4nuU29iQDtZxJT2zgyUvA77AbCWbSEjyHc5sZ8o9JMLYdTmyQ4DEgoCX9G0hLOhv/YzYJxn9Eackxk/zAcziBRwLoMMYGYrSceVMmzcBz2HCGxGNmi4Rj6cZtPDkEtBxP3yb5s+ewvHx/MuONDzXWsv3sZ/BrAQFm3h4oKwGI2QiqBwLGHz+IUTYKRsEoGAUjFgAAdiVCMY+IoHcAAAAASUVORK5CYII=","orcid":"https://orcid.org/0000-0002-5038-3535","institution":"Zhejiang Agriculture and Forestry University: Zhejiang A and F University","correspondingAuthor":true,"prefix":"","firstName":"Junhao","middleName":"","lastName":"Huang","suffix":""}],"badges":[],"createdAt":"2025-05-09 06:04:34","currentVersionCode":1,"declarations":"","doi":"10.21203/rs.3.rs-6625409/v1","doiUrl":"https://doi.org/10.21203/rs.3.rs-6625409/v1","draftVersion":[],"editorialEvents":[{"content":"https://doi.org/10.1007/s11104-025-07828-2","type":"published","date":"2025-09-03T15:57:31+00:00"}],"editorialNote":"","failedWorkflow":false,"files":[{"id":84192152,"identity":"f81b5221-19b2-46c7-bc68-459beeafea78","added_by":"auto","created_at":"2025-06-09 06:56:40","extension":"png","order_by":1,"title":"Figure 1","display":"","copyAsset":false,"role":"figure","size":205196,"visible":true,"origin":"","legend":"\u003cp\u003ePermutational multivariate analysis of variance (PERMANOVA) on the soil fauna community composition in the litter layer based on Bray-Curtis distance.\u003c/p\u003e\n\u003cp\u003eNote: *, \u003cem\u003eP\u003c/em\u003e \u0026lt; 0.05; **, \u003cem\u003eP\u003c/em\u003e \u0026lt; 0.01; ***, \u003cem\u003eP\u003c/em\u003e\u0026lt; 0.001.\u003c/p\u003e","description":"","filename":"1.png","url":"https://assets-eu.researchsquare.com/files/rs-6625409/v1/2faae8c3bd79184fcfbf8dc8.png"},{"id":84190786,"identity":"0a671861-e34b-4b31-9ec9-37a0dbc651ab","added_by":"auto","created_at":"2025-06-09 06:48:40","extension":"png","order_by":2,"title":"Figure 2","display":"","copyAsset":false,"role":"figure","size":98061,"visible":true,"origin":"","legend":"\u003cp\u003ePermutational multivariate analysis of variance (PERMANOVA) on the soil fauna community composition in the soil layer based on Bray-Curtis distance.\u003c/p\u003e\n\u003cp\u003eNote: *, \u003cem\u003eP\u003c/em\u003e \u0026lt; 0.05; **, \u003cem\u003eP\u003c/em\u003e \u0026lt; 0.01; ***, \u003cem\u003eP\u003c/em\u003e\u0026lt; 0.001.\u003c/p\u003e","description":"","filename":"2.png","url":"https://assets-eu.researchsquare.com/files/rs-6625409/v1/ae757d376f9495e173c553dc.png"},{"id":84192155,"identity":"23d42f5e-9cd9-4ea9-a4c4-008f0105e067","added_by":"auto","created_at":"2025-06-09 06:56:40","extension":"png","order_by":3,"title":"Figure 3","display":"","copyAsset":false,"role":"figure","size":71241,"visible":true,"origin":"","legend":"\u003cp\u003eRedundancy Analysis (RDA) plot showing the preference of four feeding guilds for initial litter physicochemical properties.\u003c/p\u003e\n\u003cp\u003eNate: C: carbon; N: nitrogen; TC: total carbon; TN: total nitrogen.\u003c/p\u003e","description":"","filename":"3.png","url":"https://assets-eu.researchsquare.com/files/rs-6625409/v1/8a4cbcf34ef5a1bfec32a97a.png"},{"id":84190795,"identity":"da7f40e8-7236-4bf7-874b-bfe159090baa","added_by":"auto","created_at":"2025-06-09 06:48:40","extension":"png","order_by":4,"title":"Figure 4","display":"","copyAsset":false,"role":"figure","size":204899,"visible":true,"origin":"","legend":"\u003cp\u003eTriangular plots of beta diversity comparisons (using Sørensen dissimilarity index) for soil fauna (a, c) communities and feeding guilds (b, d) among all samples. Each point represents a pair of samples. Its position is determined by a triplet of values from the similarity, replacement and richness difference matrices; each triplet sums to 1.\u003c/p\u003e","description":"","filename":"4.png","url":"https://assets-eu.researchsquare.com/files/rs-6625409/v1/50ee99896bee5ffed73d1132.png"},{"id":84190796,"identity":"fa84fe94-f1ea-4b33-9c17-ca8b16cd3bb5","added_by":"auto","created_at":"2025-06-09 06:48:40","extension":"png","order_by":5,"title":"Figure 5","display":"","copyAsset":false,"role":"figure","size":113323,"visible":true,"origin":"","legend":"\u003cp\u003eSoil fauna specific and shared OTUs under reciprocal litter transplant in litter layer (a) and soil layers (b).\u003c/p\u003e","description":"","filename":"5.png","url":"https://assets-eu.researchsquare.com/files/rs-6625409/v1/a09f6f6a51aa6c3a1ddbe08a.png"},{"id":90828107,"identity":"d5dc7719-edb9-4f7d-9ba2-e104175ee03a","added_by":"auto","created_at":"2025-09-08 16:05:48","extension":"pdf","order_by":0,"title":"","display":"","copyAsset":false,"role":"manuscript-pdf","size":1660955,"visible":true,"origin":"","legend":"","description":"","filename":"manuscript.pdf","url":"https://assets-eu.researchsquare.com/files/rs-6625409/v1/7d4245a7-b69f-4894-9120-cbc31348176c.pdf"},{"id":84190793,"identity":"5439121a-259e-441d-be24-8356d8b2ddf2","added_by":"auto","created_at":"2025-06-09 06:48:40","extension":"docx","order_by":1,"title":"","display":"","copyAsset":false,"role":"supplement","size":1440589,"visible":true,"origin":"","legend":"","description":"","filename":"Supplementalmaterials20250509.docx","url":"https://assets-eu.researchsquare.com/files/rs-6625409/v1/24a94d841e368a110199c712.docx"},{"id":84190790,"identity":"59ab9cc4-1dfe-432f-acc2-0c896811a08c","added_by":"auto","created_at":"2025-06-09 06:48:40","extension":"docx","order_by":2,"title":"","display":"","copyAsset":false,"role":"supplement","size":236977,"visible":true,"origin":"","legend":"","description":"","filename":"Table2.docx","url":"https://assets-eu.researchsquare.com/files/rs-6625409/v1/6283e244ad382aa3db035db4.docx"}],"financialInterests":"","formattedTitle":"Litter quality drives the shift of soil fauna structure without changing its ecological function","fulltext":[{"header":"1. Introduction","content":"\u003cp\u003eInvasive alien plant species are recognized as driving biodiversity loss and altering ecosystem services and socioeconomic conditions (Bacher et al., \u003cspan citationid=\"CR7\" class=\"CitationRef\"\u003e2018\u003c/span\u003e; Rai and Singh, \u003cspan citationid=\"CR73\" class=\"CitationRef\"\u003e2020\u003c/span\u003e; P\u0026eacute;rez et al., \u003cspan citationid=\"CR67\" class=\"CitationRef\"\u003e2022\u003c/span\u003e; Gallardo et al., \u003cspan citationid=\"CR34\" class=\"CitationRef\"\u003e2024\u003c/span\u003e). The ecological disturbances caused by invasive plants have been increasingly recognized as a key challenge to global sustainability (Rai and Singh, \u003cspan citationid=\"CR73\" class=\"CitationRef\"\u003e2020\u003c/span\u003e; Premakumari et al., \u003cspan citationid=\"CR70\" class=\"CitationRef\"\u003e2022\u003c/span\u003e; Khattak et al., \u003cspan citationid=\"CR42\" class=\"CitationRef\"\u003e2024\u003c/span\u003e). These species impede native plant growth by affecting soil cover, nutrient cycling, fire patterns, and hydrology (Weidlich et al., \u003cspan citationid=\"CR94\" class=\"CitationRef\"\u003e2020\u003c/span\u003e; Raheem et al., \u003cspan citationid=\"CR71\" class=\"CitationRef\"\u003e2024\u003c/span\u003e) and trigger a series of environmental effects that reshape community composition, biotic interactions (Vil\u0026agrave; et al., \u003cspan citationid=\"CR89\" class=\"CitationRef\"\u003e2011\u003c/span\u003e; David et al., \u003cspan citationid=\"CR28\" class=\"CitationRef\"\u003e2017\u003c/span\u003e; Li et al., \u003cspan citationid=\"CR43\" class=\"CitationRef\"\u003e2022\u003c/span\u003e; Zhang, \u003cspan citationid=\"CR107\" class=\"CitationRef\"\u003e2023b\u003c/span\u003e), and ecosystem processes, influencing ecosystem productivity and human well-being (Xu et al., \u003cspan citationid=\"CR99\" class=\"CitationRef\"\u003e2020\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eIn recent years, the invasion of moso bamboo (\u003cem\u003ePhyllostachys edulis\u003c/em\u003e) has emerged as an ecological concern, not only in its native region in China but also in areas where it has been introduced, such as Japan and North America (Fukushima et al., \u003cspan citationid=\"CR33\" class=\"CitationRef\"\u003e2015\u003c/span\u003e; Lieurance et al., \u003cspan citationid=\"CR48\" class=\"CitationRef\"\u003e2018\u003c/span\u003e; Xu et al., \u003cspan citationid=\"CR99\" class=\"CitationRef\"\u003e2020\u003c/span\u003e). As a native species in subtropical China, moso bamboo is the most prevalent giant timber bamboo species and is extensively cultivated (Peng et al., \u003cspan citationid=\"CR66\" class=\"CitationRef\"\u003e2013\u003c/span\u003e; Li et al., \u003cspan citationid=\"CR44\" class=\"CitationRef\"\u003e2024\u003c/span\u003e). However, in recent decades, abandoned bamboo plantations have led to the spread of moso bamboo into neighboring broadleaf forests, negatively affecting plant diversity, litter input, soil properties, and soil communities (Liu et al., \u003cspan citationid=\"CR51\" class=\"CitationRef\"\u003e2019\u003c/span\u003e; Luan et al., \u003cspan citationid=\"CR53\" class=\"CitationRef\"\u003e2021\u003c/span\u003e; Wei et al., \u003cspan citationid=\"CR93\" class=\"CitationRef\"\u003e2021\u003c/span\u003e; Li et al., \u003cspan citationid=\"CR44\" class=\"CitationRef\"\u003e2024\u003c/span\u003e). The impacts are primarily manifested as changes in litter input, root exudates, and the physical environment, facilitating invasion success by increasing primary productivity and affecting soil biota and ecosystem dynamics (Wolfe and Klironomos, \u003cspan citationid=\"CR96\" class=\"CitationRef\"\u003e2005\u003c/span\u003e). Moso bamboo invasion changes the biodiversity and community of soil fauna (Wei et al., \u003cspan citationid=\"CR93\" class=\"CitationRef\"\u003e2021\u003c/span\u003e; Long et al., \u003cspan citationid=\"CR52\" class=\"CitationRef\"\u003e2023\u003c/span\u003e; Xiao et al., \u003cspan citationid=\"CR98\" class=\"CitationRef\"\u003e2023\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eLitter from invasive plants often decompose faster than that from native plants, altering soil carbon storage and nutrient levels (Ehrenfeld, \u003cspan citationid=\"CR30\" class=\"CitationRef\"\u003e2010\u003c/span\u003e). These changes in litter quantity and quality modify habitat properties, such as soil moisture and pH, affecting soil biota performance (Kappes et al., \u003cspan citationid=\"CR40\" class=\"CitationRef\"\u003e2007\u003c/span\u003e; Zhao et al., \u003cspan citationid=\"CR108\" class=\"CitationRef\"\u003e2021\u003c/span\u003e; Morris and Blackwood, \u003cspan citationid=\"CR59\" class=\"CitationRef\"\u003e2024\u003c/span\u003e). Research has demonstrated that alterations in litter inputs from invasive plants significantly affect soil organisms, including the abundance and diversity of soil biota species, thereby influencing decomposition processes and nutrient cycling (Liao et al., \u003cspan citationid=\"CR47\" class=\"CitationRef\"\u003e2008\u003c/span\u003e; Sun et al., \u003cspan citationid=\"CR84\" class=\"CitationRef\"\u003e2023\u003c/span\u003e; Zhang, \u003cspan citationid=\"CR106\" class=\"CitationRef\"\u003e2023a\u003c/span\u003e). Previous studies have primarily focused on the impacts of microorganisms; thus, the specific effects of soil fauna remain uncertain.\u003c/p\u003e \u003cp\u003eSoil fauna significantly impact decomposition processes and serve as important biodiversity reservoirs, while their diversity and functions vary and are closely linked to different aboveground systems (Peng et al., \u003cspan citationid=\"CR65\" class=\"CitationRef\"\u003e2020\u003c/span\u003e; Wang et al., \u003cspan citationid=\"CR91\" class=\"CitationRef\"\u003e2020\u003c/span\u003e; Coleman et al., \u003cspan citationid=\"CR23\" class=\"CitationRef\"\u003e2024\u003c/span\u003e). However, there is no definitive conclusion regarding their correlation. Soil fauna are integral ecosystem components, playing crucial roles in accelerating organic matter decomposition and nutrient cycling, primarily by breaking down litter and stimulating microbial activity (Petersen, \u003cspan citationid=\"CR68\" class=\"CitationRef\"\u003e1982\u003c/span\u003e; Seastedt and Crossley Jr, \u003cspan citationid=\"CR78\" class=\"CitationRef\"\u003e1983\u003c/span\u003e; Bardgett and Chan, \u003cspan citationid=\"CR10\" class=\"CitationRef\"\u003e1999\u003c/span\u003e; Yin et al., \u003cspan citationid=\"CR104\" class=\"CitationRef\"\u003e2010\u003c/span\u003e; Reddy et al., \u003cspan citationid=\"CR74\" class=\"CitationRef\"\u003e2024\u003c/span\u003e). Soil fauna from different functional groups contribute differently to litter decomposition, with the most fundamental functional distinctions being shredders (involved in fragmentation), i.e., detritivores, and predators (Faber and Verhoef, \u003cspan citationid=\"CR31\" class=\"CitationRef\"\u003e1991\u003c/span\u003e; Brussaard, \u003cspan citationid=\"CR16\" class=\"CitationRef\"\u003e1998\u003c/span\u003e; Heděnec et al., \u003cspan citationid=\"CR37\" class=\"CitationRef\"\u003e2022\u003c/span\u003e; Coleman et al., \u003cspan citationid=\"CR23\" class=\"CitationRef\"\u003e2024\u003c/span\u003e). Shredders break down plant litter, increasing food resources for soil microfauna and mesofauna (Brown, \u003cspan citationid=\"CR15\" class=\"CitationRef\"\u003e1995\u003c/span\u003e; Olivia and Dumitru, \u003cspan citationid=\"CR63\" class=\"CitationRef\"\u003e2023\u003c/span\u003e). Fragmentation enhances the surface area of litter substrates, allowing microbes to access nutrient-rich inner tissues and thereby influencing litter decomposition (Faber and Verhoef, \u003cspan citationid=\"CR31\" class=\"CitationRef\"\u003e1991\u003c/span\u003e; Chapin et al., \u003cspan citationid=\"CR19\" class=\"CitationRef\"\u003e2002\u003c/span\u003e; Coleman et al., \u003cspan citationid=\"CR22\" class=\"CitationRef\"\u003e2017\u003c/span\u003e; Cavallet et al., \u003cspan citationid=\"CR18\" class=\"CitationRef\"\u003e2022\u003c/span\u003e). As evidenced, changes in the soil fauna community composition impact litter decomposition and nutrient metabolism (Verhoef and Brussaard, \u003cspan citationid=\"CR88\" class=\"CitationRef\"\u003e1990\u003c/span\u003e; Set\u0026auml;l\u0026auml; et al., \u003cspan citationid=\"CR79\" class=\"CitationRef\"\u003e1996\u003c/span\u003e; Jones et al., \u003cspan citationid=\"CR39\" class=\"CitationRef\"\u003e1998\u003c/span\u003e; Li et al., \u003cspan citationid=\"CR45\" class=\"CitationRef\"\u003e2021\u003c/span\u003e; Yang et al., \u003cspan citationid=\"CR100\" class=\"CitationRef\"\u003e2022\u003c/span\u003e; Balasubramanian et al., \u003cspan citationid=\"CR8\" class=\"CitationRef\"\u003e2023\u003c/span\u003e). Would changes in litter (e.g., litter quality) affect soil fauna? Peng et al. (\u003cspan citationid=\"CR64\" class=\"CitationRef\"\u003e2022\u003c/span\u003e) speculated that leaf litter quality likely serves as the primary driver for the effects of tree species on soil fauna communities. However, this speculation has not yet been confirmed.\u003c/p\u003e \u003cp\u003eLitter transplant is an effective method of litter control that confirms the \u0026ldquo;home-field advantage\u0026rdquo; (HFA) hypothesis, which states that decomposition occurs more quickly when litter decomposes beneath the plant species from which it originates (i.e., at home) than beneath different plant species (away) (Bocock et al., \u003cspan citationid=\"CR13\" class=\"CitationRef\"\u003e1960\u003c/span\u003e; Hunt et al., \u003cspan citationid=\"CR38\" class=\"CitationRef\"\u003e1988\u003c/span\u003e; Gholz et al., 2000; Negrete-Yankelevich et al., \u003cspan citationid=\"CR60\" class=\"CitationRef\"\u003e2008\u003c/span\u003e; Ayres et al., \u003cspan citationid=\"CR5\" class=\"CitationRef\"\u003e2009a\u003c/span\u003e). Previous studies have shown that litter transplant changes the microbial diversity and community structure (Milcu and Manning, \u003cspan citationid=\"CR58\" class=\"CitationRef\"\u003e2011\u003c/span\u003e; Lin et al., \u003cspan citationid=\"CR49\" class=\"CitationRef\"\u003e2019\u003c/span\u003e). Dynamic changes in microbial communities in litter are instrumental in elucidating changes in component composition and material cycling during decomposition (Ma et al., \u003cspan citationid=\"CR54\" class=\"CitationRef\"\u003e2023\u003c/span\u003e). Therefore, researchers are interested in the effects of the decomposition process on microbial communities in litter transplant experiments (Zhong et al., \u003cspan citationid=\"CR109\" class=\"CitationRef\"\u003e2018\u003c/span\u003e), while research on soil fauna is still relatively scarce.\u003c/p\u003e \u003cp\u003eThe reciprocal transplant of litter between invasive and native plants offers a potential strategy for investigating the effects of altering litter inputs on soil fauna communities. Understanding how alternations in litter quality during plant invasion affect decomposers, such as soil fauna that act as shredders, will provide further insights into ecosystem dynamics and management strategies. We focused on moso bamboo as the experimental subject to investigate the effects of litter quality on soil fauna in plant invasions. We conducted litter exchange and retention treatments between two forest stands, i.e., secondary broadleaf forest and \u003cem\u003ePhyllostachys edulis\u003c/em\u003e forest, representing two invasion stages: uninvaded and completely invaded. We investigated the impact of bamboo invasion on soil fauna diversity at both the taxonomic and functional levels as well as their community composition. For this experiment, the following scientific questions were posed: (1) Does reciprocal litter transplant (exchanging litter of different qualities) affect soil fauna diversity and community structure? (2) Does changing litter quality through reciprocal transplant affect the ecological functions performed by soil fauna?\u003c/p\u003e"},{"header":"2. Materials and methods","content":"\u003cdiv id=\"Sec3\" class=\"Section2\"\u003e \u003ch2\u003e2.1 Study site\u003c/h2\u003e \u003cp\u003eThe study site was in the Tianmu Mountain National Nature Reserve, located in Zhejiang Province, Eastern China (30\u0026deg;18'\u0026ndash;30\u0026deg;25'N, 119\u0026deg;23'\u0026ndash;119\u0026deg;29'E) at an elevation range of 500\u0026ndash;650 m. This region experiences a subtropical monsoon climate with an average annual temperature of 8.8\u0026ndash;14.8\u0026deg;C and an average annual rainfall of 1390\u0026ndash;1870 mm. The soil in the study site was derived from silt stone, classified as a Ferralsol according to the Food and Agriculture Organization of the United Nations soil classification system (WRB, \u003cspan citationid=\"CR97\" class=\"CitationRef\"\u003e2006\u003c/span\u003e). Following the abandonment of bamboo forests in the 1970s, the natural forest at this site heavily affected moso bamboo (\u003cem\u003ePhyllostachys edulis\u003c/em\u003e) encroachment (Chen et al., \u003cspan citationid=\"CR20\" class=\"CitationRef\"\u003e2021\u003c/span\u003e). At this site, we identified three distinct forest types, each representing a different stage of bamboo invasion (Wei et al., \u003cspan citationid=\"CR93\" class=\"CitationRef\"\u003e2021\u003c/span\u003e)): secondary broadleaf forest (SBF), an uninvaded native forest dominated by \u003cem\u003eCunninghamia lanceolata\u003c/em\u003e, \u003cem\u003eQuercus serrata\u003c/em\u003e var. \u003cem\u003ebrevipetiolata\u003c/em\u003e, and \u003cem\u003ePinus massoniana\u003c/em\u003e; mixed bamboo forest (MBF), a transition zone moderately invaded by moso bamboo; and \u003cem\u003ePhyllostachys edulis\u003c/em\u003e forest (PEF), which is completely invaded and dominated by moso bamboo. Our research selected SBF (uninvaded) and PEF (completely invaded) at the site for the present study. Both had similar elevations, slopes (50\u0026ndash;60%), soil textures, and bulk densities (Teng et al., \u003cspan citationid=\"CR86\" class=\"CitationRef\"\u003e2023\u003c/span\u003e).\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec4\" class=\"Section2\"\u003e \u003ch2\u003e2.2 Experimental design\u003c/h2\u003e \u003cp\u003eSix parallel transects (about 180 m in length and 50 m apart) along the encroaching path from broadleaf forest to moso bamboo forest were installed in 2016 in an area of about 4.5 ha (Wei et al., \u003cspan citationid=\"CR93\" class=\"CitationRef\"\u003e2021\u003c/span\u003e). For this study, four paired plots (20 m \u0026times; 20 m) of broadleaf forest and moso bamboo forest in the six transects were randomly selected (for four replicates). In each plot, three subplots (each 2.5 m \u0026times; 2.5 m, 10\u0026ndash;15 m apart) were established (for three treatments), resulting in 24 subplots (Fig. \u003cspan refid=\"MOESM1\" class=\"InternalRef\"\u003eS1\u003c/span\u003e). Nylon net fences, standing 0.3 m high with a mesh size of 5 mm, were installed around the perimeter of each subplot to prevent intrusion of the surrounding litterfall. Additionally, a top nylon net (2.5 m \u0026times; 2.5 m, 2 mm mesh size) was suspended 0.8 m above each subplot to intercept fresh litterfall falling into the plot (Fig. S2).\u003c/p\u003e \u003cp\u003eThese subplots were set up in February 2019. All aboveground litter within each subplot was cleared, followed by a 3-month buffer period. These three subplots in each plot were established in May 2019 and arranged randomly for three monthly treatments of the intercepted fresh litterfall: a) control group: litterfall retention\u0026mdash;fresh litterfall intercepted by the top nets from each forest stand was mixed monthly and evenly redistributed onto their own subplot; b) removal group: litterfall removal\u0026mdash;fresh intercepted litterfall was cleared monthly; and c) replacement group: litterfall replacement\u0026mdash;fresh litterfall intercepted from broadleaf or bamboo forests was exchanged and scattered onto the subplots of the other forest type (Fig. \u003cspan refid=\"MOESM1\" class=\"InternalRef\"\u003eS1\u003c/span\u003e). The litterfall in each litter trap was collected monthly and evenly added to the control and replacement treatments at an ambient rate. The average rate of litter input was 0.3 kg dry litter per m\u003csup\u003e2\u003c/sup\u003e per year for broadleaf forests and 0.33 kg dry litter for bamboo forests (Teng et al., \u003cspan citationid=\"CR86\" class=\"CitationRef\"\u003e2023\u003c/span\u003e).\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec5\" class=\"Section2\"\u003e \u003ch2\u003e2.3 Soil fauna extraction\u003c/h2\u003e \u003cp\u003eIn May 2022 (3 years after litterfall manipulation), November 2022 (3.5 years), and May 2023 (4 years), soil fauna diversity was investigated in each subplot. Litter and soil samples were collected using the three-point sampling method in each subplot. For each point, litter and humus from an area of 25 cm \u0026times; 25 cm as well as a soil column with a diameter of 5 cm and a depth of 20 cm were collected. The collected samples were labeled and transported to the laboratory on the same day. The soil fauna were then extracted using the Berlese\u0026ndash;Tullgren funnel method (2-mm diameter mesh) (Macfadyen, \u003cspan citationid=\"CR55\" class=\"CitationRef\"\u003e1953\u003c/span\u003e) in 48 h and gathered in conical flasks containing anhydrous ethanol. Soil fauna were preliminarily classified and categorized based on their morphology, as observed under a microscope (VHX-5000, Keyence Corporation, Osaka City, Japan). In each plot, soil fauna individuals from litter or soil samples from the three sampling points were bulked into one composite sample. In total, 120 samples were collected, with 72 from the soil layer (2 forest types \u0026times; 3 litter manipulations \u0026times; 4 replicates \u0026times; 3 times sampling) and 48 from the litter layer (no samples from the removal treatment). The three collected samples were considered as repeated measurements.\u003c/p\u003e \u003cp\u003eThe litterfall from the removal subplots in the two forest types was collected in November and December 2022. Since the two months encompassed the peak leaf-fall period for both bamboo and broadleaf forests, the litterfall was used to determine the initial physicochemical properties of litter inputs (the contents of cellulose, lignin, total carbon, and total nitrogen), following the protocols described by Teng et al. (\u003cspan citationid=\"CR86\" class=\"CitationRef\"\u003e2023\u003c/span\u003e) (Table \u003cspan refid=\"MOESM1\" class=\"InternalRef\"\u003eS1\u003c/span\u003e).\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec6\" class=\"Section2\"\u003e \u003ch2\u003e2.4 High-throughput sequencing (HTS) and data processing\u003c/h2\u003e \u003cp\u003eWe employed a highly efficient and reliable high-throughput sequencing (HTS) technique (Arribas et al., \u003cspan citationid=\"CR2\" class=\"CitationRef\"\u003e2016\u003c/span\u003e; Creedy et al., \u003cspan citationid=\"CR26\" class=\"CitationRef\"\u003e2019\u003c/span\u003e) to acquire the DNA sequences of the soil fauna (Wei et al., \u003cspan citationid=\"CR93\" class=\"CitationRef\"\u003e2021\u003c/span\u003e; Long et al., \u003cspan citationid=\"CR52\" class=\"CitationRef\"\u003e2023\u003c/span\u003e). All 120 samples underwent grinding and homogenization with liquid nitrogen for DNA extraction using the DNeasy\u0026reg; Blood and Tissue Kit (QIAGEN, Hilden, Germany) following the sequencing and data processing protocols outlined by Wei et al. (\u003cspan citationid=\"CR93\" class=\"CitationRef\"\u003e2021\u003c/span\u003e). Primers Ill_B_F (5\u0026prime;. CCIGAYATRGCITTYCCICG. 3\u0026prime;) (Shokralla et al., \u003cspan citationid=\"CR82\" class=\"CitationRef\"\u003e2015\u003c/span\u003e) and Fol_degen_rev (5\u0026prime;. TANACYTCNGGRTGNCCRAARAAYCA. 3\u0026prime;) (Yu et al., \u003cspan citationid=\"CR105\" class=\"CitationRef\"\u003e2012\u003c/span\u003e) were utilized in polymerase chain reaction (PCR) to amplify the 418-bp region of the cytochrome oxidase subunit I (COI) gene (barcode fragment). Each sample was amplified in triplicate on an ABI GeneAmp\u0026reg;9700 (Applied Biosystems, Waltham, Massachusetts, USA), following the PCR protocol outlined by Arribas et al. (\u003cspan citationid=\"CR2\" class=\"CitationRef\"\u003e2016\u003c/span\u003e). After the dual-index barcodes (Illumina TruSeqTM DNA Sample Prep Kit, Illumina, Inc., San Diego, California, USA) were added, the purified amplicons were pooled and sequenced on the Illumina MiSeq Platform (2 \u0026times; 300 bp paired-end) following quality control. The raw sequences underwent quality filtration using Trimmomatic (Bolger et al., \u003cspan citationid=\"CR14\" class=\"CitationRef\"\u003e2014\u003c/span\u003e) and were merged using FLASH (Magoč and Salzberg, \u003cspan citationid=\"CR56\" class=\"CitationRef\"\u003e2011\u003c/span\u003e). Only contigs with a length of 418 bp were retained.\u003c/p\u003e \u003cp\u003ePost-joining, the DNA sequences were processed and optimized using the DADA2 denoising method to obtain amplicon sequence variants (ASVs). Subsequently, TaxonDNA software (version 1.0) was employed to cluster the ASVs at a 97% threshold to generate operational taxonomic units (OTUs). To prevent sequencing errors, we removed low-abundance OTUs (total counts\u0026thinsp;\u0026lt;\u0026thinsp;5). Most soil fauna sequences generated from field biodiversity investigations could not be identified to the species level from any reference database, lacking deep coverage for soil invertebrates (Wei et al., \u003cspan citationid=\"CR93\" class=\"CitationRef\"\u003e2021\u003c/span\u003e; Long et al., \u003cspan citationid=\"CR52\" class=\"CitationRef\"\u003e2023\u003c/span\u003e). The data sorting procedure in this study fully combined the broad advantages of the two dominant marker gene data analysis methods, i.e., ASVs and OTUs, to accurately measure their diversity and applicability to communities. ASVs controlled errors sufficiently, and the resulting OTUs were regarded as the biological counterpart to a species in improved resolution (Callahan et al., \u003cspan citationid=\"CR17\" class=\"CitationRef\"\u003e2017\u003c/span\u003e). Subsequently, 1287 OTUs were annotated at the family level using the NCBI nt database, and those with an abundance greater than 0.1% were classified into 1 of 4 feeding guilds based on the literature, i.e., detritivores, herbivores, microbivores, and predators (Table S2) (Wei et al., \u003cspan citationid=\"CR93\" class=\"CitationRef\"\u003e2021\u003c/span\u003e).\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec7\" class=\"Section2\"\u003e \u003ch2\u003e2.5 Data analyses\u003c/h2\u003e \u003cp\u003eThe diversity indices (species richness, Simpson index, and Shannon\u0026ndash;Wiener) (Thukral, \u003cspan citationid=\"CR87\" class=\"CitationRef\"\u003e2017\u003c/span\u003e) and community composition (species distribution in each sample) of soil fauna were determined based on the occurrence of each OTU (=\u0026thinsp;species) in individual samples. All statistical analyses were performed using R version 4.3.2 (Team, 2013). The diversity of the soil fauna community was computed using the R packages \u0026ldquo;picante\u0026rdquo; and \u0026ldquo;vegan\u0026rdquo; (Shannon, \u003cspan citationid=\"CR80\" class=\"CitationRef\"\u003e2001\u003c/span\u003e; Oksanen et al., \u003cspan citationid=\"CR61\" class=\"CitationRef\"\u003e2013a\u003c/span\u003e). Due to the non-normal data distribution, we employed the non-parametric Kruskal\u0026ndash;Wallis one-way analysis of variance, as implemented in the R package \u0026ldquo;rstatix\u0026rdquo; (Shannon, \u003cspan citationid=\"CR80\" class=\"CitationRef\"\u003e2001\u003c/span\u003e; Oksanen et al., \u003cspan citationid=\"CR61\" class=\"CitationRef\"\u003e2013a\u003c/span\u003e) to assess the impact of reciprocal litter transplant on soil fauna diversity (Daniel, \u003cspan citationid=\"CR27\" class=\"CitationRef\"\u003e1990\u003c/span\u003e). Furthermore, we performed principal coordinate analysis (PCoA) based on the Bray\u0026ndash;Curtis distance using the R packages \u0026ldquo;vegan\u0026rdquo; and \u0026ldquo;ape\u0026rdquo; (Anderson and Willis, \u003cspan citationid=\"CR1\" class=\"CitationRef\"\u003e2003\u003c/span\u003e; Edwards et al., \u003cspan citationid=\"CR29\" class=\"CitationRef\"\u003e2015\u003c/span\u003e) to identify differences in community composition, dominant classes, and feeding guilds in soil fauna. The \u0026ldquo;ggplot2\u0026rdquo; (Wickham, \u003cspan citationid=\"CR95\" class=\"CitationRef\"\u003e2011\u003c/span\u003e) package was used to visualize the PCoA plot.\u003c/p\u003e \u003cp\u003eThe \u0026ldquo;adespatial\u0026rdquo; package (Shen et al., \u003cspan citationid=\"CR81\" class=\"CitationRef\"\u003e2020\u003c/span\u003e) was used to analyze the community and elucidate the factors driving changes in soil fauna community composition through three key processes: species replacement, richness differences, and Jaccard similarity. An UpSet plot was generated using the \u0026ldquo;UpSetR\u0026rdquo; package to depict the shared and unique OTUs among treatment groups (Conway et al., \u003cspan citationid=\"CR24\" class=\"CitationRef\"\u003e2017\u003c/span\u003e). Redundancy analysis (RDA) was performed using the \u0026ldquo;vegan\u0026rdquo; package (Oksanen et al., \u003cspan citationid=\"CR62\" class=\"CitationRef\"\u003e2013b\u003c/span\u003e) and \u0026ldquo;ggplot2\u0026rdquo; (Wickham, \u003cspan citationid=\"CR95\" class=\"CitationRef\"\u003e2011\u003c/span\u003e) to explore the correlation between soil fauna and environmental constraints (physicochemical litter properties).\u003c/p\u003e \u003c/div\u003e"},{"header":"3. Results","content":"\u003cdiv id=\"Sec9\" class=\"Section2\"\u003e\n \u003ch2\u003e3.1 Responses of soil fauna diversity to reciprocal litter transplant\u003c/h2\u003e\n \u003cp\u003eIn both bamboo and broadleaf forests, the diversity of soil fauna in both the litter and soil layer was not significantly influenced by litter retention, removal, or replacement (exchange) treatments (Fig. S3, \u003cem\u003eP\u003c/em\u003e\u0026thinsp;\u0026gt;\u0026thinsp;0.05). Similarly, in both bamboo and broadleaf forests, soil fauna diversity showed no significant response to litter treatments in the soil layer (Fig. S3, \u003cem\u003eP\u003c/em\u003e\u0026thinsp;\u0026gt;\u0026thinsp;0.05). However, significant differences were observed in the diversity indices of the soil fauna communities between the litter and the soil layer (Fig. S3, \u003cem\u003eP\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.05).\u003c/p\u003e\n\u003c/div\u003e\n\u003cdiv id=\"Sec10\" class=\"Section2\"\u003e\n \u003ch2\u003e3.2 Responses of soil fauna community composition to reciprocal litter transplant\u003c/h2\u003e\n \u003cp\u003eThe community composition of total soil fauna in both the litter and soil layer was significantly affected by reciprocal litter transplant (Table \u003cspan class=\"InternalRef\"\u003e1\u003c/span\u003e, Fig. \u003cspan class=\"InternalRef\"\u003e1\u003c/span\u003ea, P\u0026thinsp;\u0026lt;\u0026thinsp;0.001, Fig. \u003cspan class=\"InternalRef\"\u003e2\u003c/span\u003ea, P\u0026thinsp;\u0026lt;\u0026thinsp;0.01). Specifically, reciprocal litter transplant influenced the community compositions of Arachnida, Collembola, and Insecta in the litter layer, but no effect was detected in the soil layer (Table \u003cspan class=\"InternalRef\"\u003e1\u003c/span\u003e). Within the litter layer, significant distinctions were observed among all treatments regarding soil fauna communities (Table \u003cspan class=\"InternalRef\"\u003e2\u003c/span\u003ea, P\u0026thinsp;\u0026lt;\u0026thinsp;0.001; b, \u003cem\u003eP\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.001; c, \u003cem\u003eP\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.05; d, \u003cem\u003eP\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.01; e, \u003cem\u003eP\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.001).\u003c/p\u003e\n \u003ctable id=\"Tab1\" border=\"1\"\u003e\n \u003ccaption language=\"En\"\u003e\n \u003cdiv class=\"CaptionNumber\"\u003eTable 1\u003c/div\u003e\n \u003cdiv class=\"CaptionContent\"\u003e\n \u003cp\u003eResults of permutational multivariate analysis of variance (PERMANOVA) testing (F-values) the effects of reciprocal litter transplant on the community compositions of total soil fauna, dominant classes, and feeding guilds.\u003c/p\u003e\n \u003c/div\u003e\n \u003c/caption\u003e\n \u003cthead\u003e\n \u003ctr\u003e\n \u003cth align=\"left\" rowspan=\"2\"\u003e\n \u003cp\u003eGroups\u003c/p\u003e\n \u003c/th\u003e\n \u003cth align=\"left\" rowspan=\"2\"\u003e\n \u003cp\u003eTotal soil fauna\u003c/p\u003e\n \u003c/th\u003e\n \u003cth align=\"left\" colspan=\"3\"\u003e\n \u003cp\u003eDominant Classes\u003c/p\u003e\n \u003c/th\u003e\n \u003cth align=\"left\"\u003e\u0026nbsp;\u003c/th\u003e\n \u003cth align=\"left\" colspan=\"4\"\u003e\n \u003cp\u003eFeeding guilds\u003c/p\u003e\n \u003c/th\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003cth align=\"left\"\u003e\n \u003cp\u003eArachnida\u003c/p\u003e\n \u003c/th\u003e\n \u003cth align=\"left\"\u003e\n \u003cp\u003eCollembola\u003c/p\u003e\n \u003c/th\u003e\n \u003cth align=\"left\"\u003e\n \u003cp\u003eInsecta\u003c/p\u003e\n \u003c/th\u003e\n \u003cth align=\"left\"\u003e\u0026nbsp;\u003c/th\u003e\n \u003cth align=\"left\"\u003e\n \u003cp\u003eDetritivores\u003c/p\u003e\n \u003c/th\u003e\n \u003cth align=\"left\"\u003e\n \u003cp\u003eHerbivores\u003c/p\u003e\n \u003c/th\u003e\n \u003cth align=\"left\"\u003e\n \u003cp\u003eMicrobivores\u003c/p\u003e\n \u003c/th\u003e\n \u003cth align=\"left\"\u003e\n \u003cp\u003ePredators\u003c/p\u003e\n \u003c/th\u003e\n \u003c/tr\u003e\n \u003c/thead\u003e\n \u003ctbody\u003e\n \u003ctr\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e\u003cstrong\u003eLitter layer\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e0.0913***\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e0.1021***\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e0.0981***\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e0.0788***\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\u0026nbsp;\u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e0.1372**\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e0.1149**\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e0.1182**\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e0.1174***\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e\u003cstrong\u003eSoil layer\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e0.0793**\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e0.0927\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e0.1275\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e0.0745\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\u0026nbsp;\u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e0.1031\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e0.1284\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e0.0713\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e0.1081**\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003c/tbody\u003e\n \u003ctfoot\u003e\n \u003ctr\u003e\n \u003ctd colspan=\"10\"\u003eNote: *, \u003cem\u003eP\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.05; **, \u003cem\u003eP\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.01; ***, \u003cem\u003eP\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.001.\u003c/td\u003e\n \u003c/tr\u003e\n \u003c/tfoot\u003e\n \u003c/table\u003e\n \u003cp\u003eHowever, the results varied in the soil layer. The soil fauna communities in the bamboo forest did not show significant differences under two litter types (Table \u003cspan class=\"InternalRef\"\u003e2\u003c/span\u003eh, P\u0026thinsp;\u0026gt;\u0026thinsp;0.05), whereas a difference was observed in the broadleaf forest (Table \u003cspan class=\"InternalRef\"\u003e2\u003c/span\u003eg, P\u0026thinsp;\u0026lt;\u0026thinsp;0.01). Between the two forest stands, bamboo litter significantly influenced the soil fauna community composition (Table \u003cspan class=\"InternalRef\"\u003e2\u003c/span\u003ei, P\u0026thinsp;\u0026gt;\u0026thinsp;0.05; j, \u003cem\u003eP\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.01). Furthermore, in the bamboo forest, the soil fauna community composition was not significantly affected by the removal, retention, or replacement of litter (Table \u003cspan class=\"InternalRef\"\u003e2\u003c/span\u003eh, P\u0026thinsp;\u0026gt;\u0026thinsp;0.05; l, \u003cem\u003eP\u003c/em\u003e\u0026thinsp;\u0026gt;\u0026thinsp;0.05; n, \u003cem\u003eP\u003c/em\u003e\u0026thinsp;\u0026gt;\u0026thinsp;0.05). In contrast, in the broadleaf forest, bamboo litter had a significant impact on the soil fauna community structure (Table \u003cspan class=\"InternalRef\"\u003e2\u003c/span\u003eg, P\u0026thinsp;\u0026lt;\u0026thinsp;0.01; k, \u003cem\u003eP\u003c/em\u003e\u0026thinsp;\u0026gt;\u0026thinsp;0.05; m, \u003cem\u003eP\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.01).\u003c/p\u003e\n\u003c/div\u003e\n\u003cdiv id=\"Sec11\" class=\"Section2\"\u003e\n \u003ch2\u003e3.3 Responses of soil fauna feeding guilds to reciprocal litter transplant\u003c/h2\u003e\n \u003cp\u003eAcross the four feeding guilds, soil fauna within the soil layer demonstrated lower susceptibility to reciprocal litter transplant compared to those in the litter layer since no effects were observed on herbivores, microbivores, or detritivores in the soil layer (Table \u003cspan class=\"InternalRef\"\u003e1\u003c/span\u003e, P\u0026thinsp;\u0026gt;\u0026thinsp;0.05). However, predators were notably affected by reciprocal litter transplant in both the litter and the soil layer (Table \u003cspan class=\"InternalRef\"\u003e1\u003c/span\u003e, P\u0026thinsp;\u0026lt;\u0026thinsp;0.01; Fig. \u003cspan class=\"InternalRef\"\u003e1\u003c/span\u003eh, P\u0026thinsp;\u0026lt;\u0026thinsp;0.001; Fig. \u003cspan class=\"InternalRef\"\u003e2\u003c/span\u003eb, P\u0026thinsp;\u0026lt;\u0026thinsp;0.001).\u003c/p\u003e\n \u003cp\u003eRDA revealed that the feeding guilds of soil fauna within the litter layer had varied correlations with the physicochemical properties of litter (Fig. \u003cspan class=\"InternalRef\"\u003e3\u003c/span\u003e). Herbivores exhibited a positive correlation with litter lignin and total nitrogen contents but a negative correlation with the cellulose content. Microbivores showed a positive correlation with the litter cellulose content and the carbon-to-nitrogen ratio. Predators and detritivores demonstrated a positive correlation with the total carbon content.\u003c/p\u003e\n\u003c/div\u003e\n\u003cdiv id=\"Sec12\" class=\"Section2\"\u003e\n \u003ch2\u003e3.4 Species replacement occurred in soil fauna community composition\u003c/h2\u003e\n \u003cp\u003eThe ternary plots demonstrated that the total soil fauna communities primarily underwent species replacement (Fig. \u003cspan class=\"InternalRef\"\u003e4\u003c/span\u003ea and c) with reciprocal litter transplant, but such a distinct replacement process was not detected in the feeding guild communities (Fig. \u003cspan class=\"InternalRef\"\u003e4\u003c/span\u003eb and d). This result was consistent with that from the UpSet plot (Fig. \u003cspan class=\"InternalRef\"\u003e5\u003c/span\u003e), which showed a pattern of high species exclusivity and low species sharing in the overall soil fauna community. Beta diversity decomposition analyses showed that the soil fauna community composition dissimilarities within the litter and the soil layer were primarily driven by species replacement processes, contributing 79.73 and 77.62%, respectively. In contrast, richness difference processes contributed only 20.27 and 22.38%, respectively, on average (Fig. \u003cspan class=\"InternalRef\"\u003e4\u003c/span\u003ea and c, Table S3).\u003c/p\u003e\n\u003c/div\u003e"},{"header":"4. Discussion","content":"\u003cp\u003eThis study uncovered how bamboo invasion affected soil fauna communities by altering litter quality, thus impacting the ecosystem structure, and provided direct evidence of the close link between soil fauna and aboveground systems. It also showed that despite species turnover in soil fauna, ecological functions, such as decomposition, remained stable, highlighting ecosystem resilience and informing strategies to manage invasive plants without disrupting key ecological processes.\u003c/p\u003e \u003cp\u003eWe investigated the impact of reciprocal litter transplant on soil fauna diversity and community composition in the context of bamboo invasion, addressing three scientific questions outlined in the experimental design: (1) Litter reciprocal transplant did not significantly affect soil fauna diversity but did alter community composition; (2) changes in soil fauna communities occurred through species replacement without affecting their ecological functions; and (3) changes in soil fauna communities were driven by higher litter quality.\u003c/p\u003e \u003cdiv id=\"Sec14\" class=\"Section2\"\u003e \u003ch2\u003e4.1 Litter removal did not affect soil fauna community composition\u003c/h2\u003e \u003cp\u003eThis study demonstrated that soil fauna diversity and community composition in the soil layer did not exhibit significant changes after litter removal compared to litter retention (Table\u0026nbsp;\u003cspan refid=\"Tab2\" class=\"InternalRef\"\u003e2\u003c/span\u003ek and n). Similarly, Ashford et al. (\u003cspan citationid=\"CR3\" class=\"CitationRef\"\u003e2013\u003c/span\u003e) found that litter removal did not alter soil fauna diversity or community biomass. The resilience of soil fauna may be attributed to the communities residing in deeper soil layers being less influenced by external environmental factors (Scherber et al., \u003cspan citationid=\"CR77\" class=\"CitationRef\"\u003e2010\u003c/span\u003e; Yin et al., \u003cspan citationid=\"CR103\" class=\"CitationRef\"\u003e2019\u003c/span\u003e). Long et al. (\u003cspan citationid=\"CR52\" class=\"CitationRef\"\u003e2023\u003c/span\u003e) found that surface-dwelling collembola were more susceptible to external environmental influences than their soil-dwelling counterparts. In line with this report, our results showed that the soil fauna community structure in the soil layer was less sensitive to reciprocal litter transplant than that in the litter layer (Table\u0026nbsp;\u003cspan refid=\"Tab1\" class=\"InternalRef\"\u003e1\u003c/span\u003e). In other words, local litter did not affect the soil fauna community structure in the soil layer.\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec15\" class=\"Section2\"\u003e \u003ch2\u003e4.2 The exchange of litter inputs changed soil fauna community composition\u003c/h2\u003e \u003cp\u003eThis study revealed that reciprocal litter transplant did not affect soil fauna diversity in the context of bamboo invasion (Fig. S3). This finding aligns with previous controlled experiments, which showed no significant differences in soil fauna richness and density across three forest types in a litter decomposition study (Yang and Chen, \u003cspan citationid=\"CR101\" class=\"CitationRef\"\u003e2009\u003c/span\u003e). Furthermore, Ashford et al. (\u003cspan citationid=\"CR3\" class=\"CitationRef\"\u003e2013\u003c/span\u003e) demonstrated that neither the removal nor the addition of leaf litter significantly affected taxonomic richness or diversity. However, observational investigations have shown that litter inputs from different forest types have significant effects on soil fauna richness and abundance (Cifuentes-Croquevielle et al., \u003cspan citationid=\"CR21\" class=\"CitationRef\"\u003e2020\u003c/span\u003e; Wei et al., \u003cspan citationid=\"CR93\" class=\"CitationRef\"\u003e2021\u003c/span\u003e). These contrasting results may be attributed to the varied conditions between the experiments and the natural forests, highlighting the complexity of ecological interactions in forest ecosystems. The lack of significant effects of reciprocal litter transplant on soil fauna diversity could be explained by ecological functional redundancy.\u003c/p\u003e \u003cp\u003eHowever, soil fauna community composition showed significant alterations after reciprocal litter transplant (Table\u0026nbsp;\u003cspan refid=\"Tab1\" class=\"InternalRef\"\u003e1\u003c/span\u003e). Notably, in the litter layer, significant differences were observed in soil fauna communities between litter retention and replacement treatments (Table\u0026nbsp;\u003cspan refid=\"Tab2\" class=\"InternalRef\"\u003e2\u003c/span\u003ea\u0026ndash;e), indicating the adaptation of local soil communities to the specific litter type in the decomposition process (Bocock et al., \u003cspan citationid=\"CR13\" class=\"CitationRef\"\u003e1960\u003c/span\u003e; Wardle, \u003cspan citationid=\"CR92\" class=\"CitationRef\"\u003e2002\u003c/span\u003e; Ayres et al., \u003cspan citationid=\"CR6\" class=\"CitationRef\"\u003e2009b\u003c/span\u003e; Barantal et al., \u003cspan citationid=\"CR9\" class=\"CitationRef\"\u003e2011\u003c/span\u003e; Austin et al., \u003cspan citationid=\"CR4\" class=\"CitationRef\"\u003e2014\u003c/span\u003e; Zhu et al., \u003cspan citationid=\"CR111\" class=\"CitationRef\"\u003e2024\u003c/span\u003e). The attributes of leaf litter and the forest floor, which act as key food sources and habitats, may affect the composition of soil fauna communities both directly, through providing food, and indirectly, by influencing the soil environment as a habitat (Peng et al., \u003cspan citationid=\"CR64\" class=\"CitationRef\"\u003e2022\u003c/span\u003e). Studies have consistently demonstrated that soil fauna community composition varies across regions dominated by different plant species (Griffiths et al., \u003cspan citationid=\"CR36\" class=\"CitationRef\"\u003e1992\u003c/span\u003e; Grayston et al., \u003cspan citationid=\"CR35\" class=\"CitationRef\"\u003e1998\u003c/span\u003e; Bardgett and Walker, \u003cspan citationid=\"CR11\" class=\"CitationRef\"\u003e2004\u003c/span\u003e; Wei et al., \u003cspan citationid=\"CR93\" class=\"CitationRef\"\u003e2021\u003c/span\u003e). The soil fauna community composition for decomposing the same litter type differed significantly among forest stands (Table\u0026nbsp;\u003cspan refid=\"Tab2\" class=\"InternalRef\"\u003e2\u003c/span\u003ed and e). This is likely due to community variations in the native soil fauna for each local stand (Wei et al., \u003cspan citationid=\"CR93\" class=\"CitationRef\"\u003e2021\u003c/span\u003e), with those fauna capable of successful decomposition adopting the specific manipulated litter in their forest stands. Similarly, soil fauna communities within the same forest stand showed variations among litter types (Table\u0026nbsp;\u003cspan refid=\"Tab2\" class=\"InternalRef\"\u003e2\u003c/span\u003eb and c), which may be attributed to specific litter types requiring decomposer communities with particular breakdown functions (Hunt et al., \u003cspan citationid=\"CR38\" class=\"CitationRef\"\u003e1988\u003c/span\u003e). Moreover, Li et al. (\u003cspan citationid=\"CR46\" class=\"CitationRef\"\u003e2020\u003c/span\u003e) found that soil microorganisms specialized in decomposing specific resources at different litter decomposition stages. Such specificity in adaptation allows soil organisms to effectively utilize the litter from particular plant species, which may, in turn, influence organic material cycling and the dynamic balance within soil ecosystem functions.\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec16\" class=\"Section2\"\u003e \u003ch2\u003e4.3 Litter quality drives changes in soil fauna communities\u003c/h2\u003e \u003cp\u003eIn this study\u0026rsquo;s pairwise comparison of three reciprocal litter replacement manipulations, a significant difference in soil-dwelling fauna was only detected when bamboo litter was placed in the broadleaf forest (Table\u0026nbsp;\u003cspan refid=\"Tab2\" class=\"InternalRef\"\u003e2\u003c/span\u003e). Moreover, in the litter layer, the influence of bamboo litter on soil fauna communities was more pronounced than that of broadleaf litter (Table\u0026nbsp;\u003cspan refid=\"Tab2\" class=\"InternalRef\"\u003e2\u003c/span\u003e). This may be related to the high quality of bamboo litter, as it has an elevated N concentration and low lignin/N ratio (Table \u003cspan refid=\"MOESM1\" class=\"InternalRef\"\u003eS1\u003c/span\u003e) (Phillips et al., \u003cspan citationid=\"CR69\" class=\"CitationRef\"\u003e2013\u003c/span\u003e). Studies have shown that high-quality litter (e.g., high N content and low lignin, phenol, and tannin contents) (Rai et al., \u003cspan citationid=\"CR72\" class=\"CitationRef\"\u003e2016\u003c/span\u003e; Lin et al., \u003cspan citationid=\"CR49\" class=\"CitationRef\"\u003e2019\u003c/span\u003e) ensure greater availability of essential nutrients, such as phosphorus, calcium, and magnesium, in the forest floor, which are critical factors influencing soil fauna communities (Yang et al., \u003cspan citationid=\"CR102\" class=\"CitationRef\"\u003e2020\u003c/span\u003e; Peng et al., \u003cspan citationid=\"CR64\" class=\"CitationRef\"\u003e2022\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eSpecifically, high-quality litter are harder to decompose (Melillo et al., \u003cspan citationid=\"CR57\" class=\"CitationRef\"\u003e1982\u003c/span\u003e; Vivanco and Austin, \u003cspan citationid=\"CR90\" class=\"CitationRef\"\u003e2008\u003c/span\u003e; Coq et al., \u003cspan citationid=\"CR25\" class=\"CitationRef\"\u003e2010\u003c/span\u003e) and may stimulate native decomposer communities more effectively. In low-quality broadleaf forests, the presence of higher quality bamboo litter significantly altered the soil fauna community composition. In other words, high-quality litter input had a greater impact on the soil fauna community in low-quality litter environments but not vice versa. This finding confirms that leaf litter quality is likely the primary factor behind the effect of tree species on the soil fauna community (Peng et al., \u003cspan citationid=\"CR64\" class=\"CitationRef\"\u003e2022\u003c/span\u003e). The results align with those of previous studies indicating that the decomposer community's capacity to adapt to varying litter qualities may be limited by its adaptation to locally prevalent, low-quality litter (Austin et al., \u003cspan citationid=\"CR4\" class=\"CitationRef\"\u003e2014\u003c/span\u003e; Zhou et al., \u003cspan citationid=\"CR110\" class=\"CitationRef\"\u003e2020\u003c/span\u003e). This phenomenon may be due to the enhanced decomposition rate of bamboo litter, which is facilitated by shifts in soil biota and its higher quality compared to native tree species, thus aiding the invasion process (Luan et al., \u003cspan citationid=\"CR53\" class=\"CitationRef\"\u003e2021\u003c/span\u003e; Liu et al., \u003cspan citationid=\"CR50\" class=\"CitationRef\"\u003e2024\u003c/span\u003e). This could be a successful strategy for bamboo invasion.\u003c/p\u003e \u003cp\u003e \u003cb\u003e4.4 Reciprocal litter transplant altered the soil fauna community through species replacement without changing its ecological function\u003c/b\u003e \u003c/p\u003e \u003cp\u003eThis study found that alterations in soil fauna community composition were primarily driven by the species replacement process, which is consistently observed both in the litter and the soil layer (Fig.\u0026nbsp;\u003cspan refid=\"Fig4\" class=\"InternalRef\"\u003e4\u003c/span\u003ea and c). This aligns with the predictions of Filgueiras et al. (\u003cspan citationid=\"CR32\" class=\"CitationRef\"\u003e2021\u003c/span\u003e), who suggested that ecological groups are predictably replaced as land-use intensifies, which significantly alters habitat availability and quality. Additionally, species replacement can be attributed to soil biological communities associated with specific plant species that specialize in decomposing litter from those plants (Bocock et al., \u003cspan citationid=\"CR13\" class=\"CitationRef\"\u003e1960\u003c/span\u003e; Ayres et al., \u003cspan citationid=\"CR6\" class=\"CitationRef\"\u003e2009b\u003c/span\u003e; Strickland et al., \u003cspan citationid=\"CR83\" class=\"CitationRef\"\u003e2009\u003c/span\u003e; Li et al., \u003cspan citationid=\"CR46\" class=\"CitationRef\"\u003e2020\u003c/span\u003e). Consistent with previous research, the high silicon content of bamboo fosters the development of specific decomposer communities during the decomposition process (Schaller et al., \u003cspan citationid=\"CR76\" class=\"CitationRef\"\u003e2016\u003c/span\u003e). Consequently, the litter input changes in our study might lead to corresponding alternations in the soil fauna associated with them.\u003c/p\u003e \u003cp\u003eHowever, the feeding guild communities (trophic levels) did not show a replacement process (Fig.\u0026nbsp;\u003cspan refid=\"Fig4\" class=\"InternalRef\"\u003e4\u003c/span\u003eb and d), indicating the stability of the forest soil trophic structure with reciprocal litter transplant. Soil fauna diversity varied greatly between native bamboo forests and broadleaf forests in subtropical regions (Wei et al., \u003cspan citationid=\"CR93\" class=\"CitationRef\"\u003e2021\u003c/span\u003e; Long et al., \u003cspan citationid=\"CR52\" class=\"CitationRef\"\u003e2023\u003c/span\u003e), potentially influencing their capacity for litter decomposition. The ability of soil fauna to decompose litter is not solely determined by their diversity but also by litter quality. When litter decompose in a new environment, litter quality changes alter the associated food web. Although the necessary soil fauna species may be absent, other species with similar feeding habits can fulfill this role, demonstrating ecological functional redundancy, which helps maintain ecosystem stability (Biggs et al., \u003cspan citationid=\"CR12\" class=\"CitationRef\"\u003e2020\u003c/span\u003e). In our study, among treatment groups, the OTUs that were replaced still belonged to the same feeding guild (Table S4). These findings align with those of Sauvadet et al. (\u003cspan citationid=\"CR75\" class=\"CitationRef\"\u003e2017\u003c/span\u003e), who found that litters with distinct qualities primarily influenced the first decomposer communities without significantly affecting indirect trophic links. In summary, species replacement in soil fauna communities primarily involves substitution with species in the same ecological niche, maintaining ecosystem stability.\u003c/p\u003e \u003c/div\u003e"},{"header":"5. Conclusion","content":"\u003cp\u003eThis study highlights how plant invasion influences the soil fauna community through alterations in litter quality. Specifically, reciprocal litter transplant during bamboo invasions reshaped soil fauna communities through species replacement. Notably, introducing high-quality litter into low-quality litter stands triggered substantial changes in the soil fauna community. Despite these shifts, the trophic structure of the soil fauna remained stable, reflecting the resilience and functional stability of local soil ecosystems. This further confirms the strong relationship between soil fauna and aboveground systems. These insights provide a potential strategy for managing plant invasions by modifying litter inputs to influence soil fauna communities. Such an approach can suppress invasive plants and preserve essential ecological functions, such as organic matter decomposition and nutrient cycling. The complex relationship between soil fauna and litter quality plays a pivotal role in maintaining ecosystem health and offers promising avenues for ecosystem management.\u003c/p\u003e "},{"header":"Declarations","content":"\u003cp\u003e\u003cstrong\u003eCRediT authorship contribution statement\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eKui Long\u003c/strong\u003e: Data curation, Formal analysis, Methodology, Software, Validation, Visualization, Writing \u0026ndash; original draft, Writing \u0026ndash; review \u0026amp; editing. \u003cstrong\u003eZhenyu Zhou\u003c/strong\u003e: Data curation, Formal analysis, Investigation, Validation, Visualization, Writing \u0026ndash; original draft. \u003cstrong\u003eGan Yu\u003c/strong\u003e: Data curation, Investigation, Formal analysis. \u003cstrong\u003eYakun Zhang\u003c/strong\u003e: Writing \u0026ndash; review \u0026amp; editing. \u003cstrong\u003eYang Wang\u003c/strong\u003e: Investigation, Formal analysis. \u003cstrong\u003eQingyun Wang\u003c/strong\u003e: Formal analysis, Project administration, Software. \u003cstrong\u003eYongchun Li\u003c/strong\u003e: Conceptualization, Funding acquisition, Project administration, Resources, Writing \u0026ndash; review \u0026amp; editing. \u003cstrong\u003eJunhao Huang\u003c/strong\u003e: Conceptualization, Funding acquisition, Methodology, Project administration, Resources, Supervision, Writing \u0026ndash; review \u0026amp; editing. All authors validated the findings and approved the final version for publication.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eData availability\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eIllumina sequence data supporting the results of this study will be submitted to NCBI once paper is accepted by the journal.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eDeclaration of interests\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eThe authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eAcknowledgements\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eThe authors are very grateful to Professor Feng Zhang (Nanjing Agricultural University), Dr. Xiaolong Lin (Shanghai Ocean University), and Professor Jianjun Guo (Guizhou University) for their great support for soil fauna identifications. The authors also thank Dr. Bismillah Shah, Ms. Yi Zhu, Mr. Ao Li, and Mr. Yisa Shao (Zhejiang A\u0026amp;F University, Hangzhou, China) for their assistance with fieldwork. This study was supported by the National Natural Science Foundation of China (NSFC, Grant No. 32071742).\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eDeclaration of generative AI and AI-assisted technologies in the writing process\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eDuring the preparation of this work the authors used ChatGPT in order to improve the language and readability of the manuscript. After using this tool, the authors reviewed and edited the content as needed and take full responsibility for the content of the publication.\u003c/p\u003e"},{"header":"References","content":"\u003col\u003e\n\u003cli\u003eAnderson, M.J., Willis, T.J., 2003. Canonical analysis of principal coordinates: a useful method of constrained ordination for ecology. Ecology 84, 511-525. https://doi.org/10.1890/0012-9658(2003)084[0511:CAOPCA]2.0.CO;2\u003c/li\u003e\n\u003cli\u003eArribas, P., And\u0026uacute;jar, C., Hopkins, K., Shepherd, M., Vogler, A.P., 2016. Metabarcoding and mitochondrial metagenomics of endogean arthropods to unveil the mesofauna of the soil. Methods Ecol. Evol. 7, 1071-1081.https://doi.org/10.1111/2041-210X.12557\u003c/li\u003e\n\u003cli\u003eAshford, O., Foster, W., Turner, B.L., Sayer, E., Sutcliffe, L., Tanner, E., 2013. Litter manipulation and the soil arthropod community in a lowland tropical rainforest. Soil Biol. Biochem. 62, 5-12. https://doi.org/10.1016/j.soilbio.2013.03.001\u003c/li\u003e\n\u003cli\u003eAustin, A.T., Vivanco, L., Gonz\u0026aacute;lez‐Arzac, A., P\u0026eacute;rez, L.I., 2014. There\u0026apos;s no place like home? An exploration of the mechanisms behind plant litter\u0026ndash;decomposer affinity in terrestrial ecosystems. Wiley Online Library.https://doi.org/10.1111/nph.12959\u003c/li\u003e\n\u003cli\u003eAyres, E., Steltzer, H., Berg, S., Wall, D.H., 2009a. Soil biota accelerate decomposition in high‐elevation forests by specializing in the breakdown of litter produced by the plant species above them. J. Ecol. 97, 901-912. https://doi.org/10.1111/j.1365-2745.2009.01539.x\u003c/li\u003e\n\u003cli\u003eAyres, E., Steltzer, H., Berg, S., Wallenstein, M.D., Simmons, B.L., Wall, D.H., 2009b. Tree species traits influence soil physical, chemical, and biological properties in high elevation forests. PloS one 4, e5964. https://doi.org/10.1371/journal.pone.0005964\u003c/li\u003e\n\u003cli\u003eBacher, S., Blackburn, T.M., Essl, F., Genovesi, P., Heikkil\u0026auml;, J., Jeschke, J.M., Jones, G., Keller, R., Kenis, M., Kueffer, C., 2018. Socio‐economic impact classification of alien taxa (SEICAT). Methods Ecol. Evol. 9, 159-168. https://doi.org/10.1111/2041-210X.12844\u003c/li\u003e\n\u003cli\u003eBalasubramanian, A., Sivakumar, B., CN, H.P., Swathiga, G., Vasanth, V., Thirumoorthy, P., Bora, N.R., SJ, M.P., 2023. Influence of Soil Invertebrates on Soil Decomposition. J. Surv. Fish. Sci. 10, 618-629. https://doi.org/10.53555/sfs.v10i3.1815\u003c/li\u003e\n\u003cli\u003eBarantal, S., Roy, J., Fromin, N., Schimann, H., H\u0026auml;ttenschwiler, S., 2011. Long-term presence of tree species but not chemical diversity affect litter mixture effects on decomposition in a neotropical rainforest. Oecologia 167, 241-252. https://doi.org/10.1007/s00442-011-1966-4\u003c/li\u003e\n\u003cli\u003eBardgett, R.D., Chan, K.F., 1999. Experimental evidence that soil fauna enhance nutrient mineralization and plant nutrient uptake in montane grassland ecosystems. Soil Biol. Biochem. 31, 1007-1014. https://doi.org/10.1016/S0038-0717(99)00014-0\u003c/li\u003e\n\u003cli\u003eBardgett, R.D., Walker, L.R., 2004. Impact of coloniser plant species on the development of decomposer microbial communities following deglaciation. Soil Biol. Biochem. 36, 555-559. https://doi.org/10.1016/j.soilbio.2003.11.002\u003c/li\u003e\n\u003cli\u003eBiggs, C.R., Yeager, L.A., Bolser, D.G., Bonsell, C., Dichiera, A.M., Hou, Z., Keyser, S.R., Khursigara, A.J., Lu, K., Muth, A.F., 2020. Does functional redundancy affect ecological stability and resilience? A review and meta-analysis. Ecosphere 11, e03184. https://doi.org/10.1002/ecs2.3184\u003c/li\u003e\n\u003cli\u003eBocock, K., Gilbert, O., Capstick, C., Twinn, D., Waid, J., Woodman, M., 1960. Changes in leaf litter when placed on the surface of soils with contrasting humus types: I. Losses in dry weight of oak and ash leaf litter. J. Soil Sci. 11, 1-9. https://doi.org/10.1111/j.1365-2389.1960.tb02196.x\u003c/li\u003e\n\u003cli\u003eBolger, A.M., Lohse, M., Usadel, B., 2014. Trimmomatic: a flexible trimmer for Illumina sequence data. Bioinformatics 30, 2114-2120. 10.1093/bioinformatics/btu170\u003c/li\u003e\n\u003cli\u003eBrown, G.G., 1995. How do earthworms affect microfloral and faunal community diversity?, The Significance and Regulation of Soil Biodiversity: Proceedings of the International Symposium on Soil Biodiversity, held at Michigan State University, East Lansing, May 3\u0026ndash;6, 1993. Springer, pp. 247-269. https://doi.org/10.1007/BF02183068\u003c/li\u003e\n\u003cli\u003eBrussaard, L., 1998. Soil fauna, guilds, functional groups and ecosystem processes. Appl. Soil Ecol. 9, 123-135. https://doi.org/10.1016/S0929-1393(98)00066-3\u003c/li\u003e\n\u003cli\u003eCallahan, B.J., McMurdie, P.J., Holmes, S.P., 2017. Exact sequence variants should replace operational taxonomic units in marker-gene data analysis. ISME J. 11, 2639-2643. https://doi.org/10.1038/ismej.2017.119\u003c/li\u003e\n\u003cli\u003eCavallet, B.V., Silva, E.R., Baretta, C.R.D.M., Rezende, R.d.S., 2022. Effect of agriculture land use on standard cellulosic substrates breakdown and invertebrates\u0026rsquo; community. Community Ecol. 23, 277-288. https://doi.org/10.1007/s42974-022-00103-9\u003c/li\u003e\n\u003cli\u003eChapin, F.S., Matson, P.A., Mooney, H.A., Vitousek, P.M., 2002. Principles of terrestrial ecosystem ecology. https://doi.org/10.1007/b97397\u003c/li\u003e\n\u003cli\u003eChen, Z., Li, Y., Chang, S.X., Xu, Q., Li, Y., Ma, Z., Qin, H., Cai, Y., 2021. Linking enhanced soil nitrogen mineralization to increased fungal decomposition capacity with Moso bamboo invasion of broadleaf forests. Sci. Total Environ. 771, 144779. https://doi.org/10.1016/j.scitotenv.2020.144779\u003c/li\u003e\n\u003cli\u003eCifuentes-Croquevielle, C., Stanton, D.E., Armesto, J.J., 2020. Soil invertebrate diversity loss and functional changes in temperate forest soils replaced by exotic pine plantations. Sci. Rep. 10, 7762. https://doi.org/10.1038/s41598-020-64453-y\u003c/li\u003e\n\u003cli\u003eColeman, D.C., Callaham, M.A., Crossley Jr, D., 2017. Fundamentals of soil ecology. Academic press. https://doi.org/10.1016/C2015-0-04083-7\u003c/li\u003e\n\u003cli\u003eColeman, D.C., Geisen, S., Wall, D., 2024. Soil fauna: Occurrence, biodiversity, and roles in ecosystem function, Soil microbiology, ecology and biochemistry. Elsevier, pp. 131-159. https://doi.org/10.1016/B978-0-12-822941-5.00005-3\u003c/li\u003e\n\u003cli\u003eConway, J.R., Lex, A., Gehlenborg, N., 2017. UpSetR: an R package for the visualization of intersecting sets and their properties. Bioinformatics 33, 2938-2940. https://doi.org/10.1093/bioinformatics/btx364\u003c/li\u003e\n\u003cli\u003eCoq, S., Souquet, J.-M., Meudec, E., Cheynier, V., H\u0026auml;ttenschwiler, S., 2010. Interspecific variation in leaf litter tannins drives decomposition in a tropical rain forest of French Guiana. Ecology 91, 2080-2091. https://doi.org/10.1890/09-1076.1\u003c/li\u003e\n\u003cli\u003eCreedy, T.J., Ng, W.S., Vogler, A.P., 2019. Toward accurate species‐level metabarcoding of arthropod communities from the tropical forest canopy. Ecol. Evol. 9, 3105-3116. https://doi.org/10.1002/ece3.4839\u003c/li\u003e\n\u003cli\u003eDaniel, W.W., 1990. Kruskal\u0026ndash;Wallis one-way analysis of variance by ranks. Applied nonparametric statistics, 226-234.\u003c/li\u003e\n\u003cli\u003eDavid, P., Thebault, E., Anneville, O., Duyck, P.-F., Chapuis, E., Loeuille, N., 2017. Impacts of invasive species on food webs: a review of empirical data. Adv. Ecol. Res. 56, 1-60. https://doi.org/10.1016/bs.aecr.2016.10.001\u003c/li\u003e\n\u003cli\u003eEdwards, J., Johnson, C., Santos-Medell\u0026iacute;n, C., Lurie, E., Podishetty, N.K., Bhatnagar, S., Eisen, J.A., Sundaresan, V., 2015. Structure, variation, and assembly of the root-associated microbiomes of rice. Proc. Natl. Acad. Sci. U.S.A. 112, E911-E920. https://doi.org/10.1073/pnas.1414592112\u003c/li\u003e\n\u003cli\u003eEhrenfeld, J.G., 2010. Ecosystem consequences of biological invasions. Annu. Rev. Ecol. Evol. Syst. 41, 59-80. https://doi.org/10.1146/annurev-ecolsys-102209-144650\u003c/li\u003e\n\u003cli\u003eFaber, J.H., Verhoef, H.A., 1991. Functional differences between closely-related soil arthropods with respect to decomposition processes in the presence or absence of pine tree roots. Soil Biol. Biochem. 23, 15-23. https://doi.org/10.1016/0038-0717(91)90157-F\u003c/li\u003e\n\u003cli\u003eFilgueiras, B.K., Peres, C.A., Melo, F.P., Leal, I.R., Tabarelli, M., 2021. Winner\u0026ndash;loser species replacements in human-modified landscapes. Trends Ecol. Evol. 36, 545-555. https://doi.org/10.1016/j.tree.2021.02.006\u003c/li\u003e\n\u003cli\u003eFukushima, K., Usui, N., Ogawa, R., Tokuchi, N., 2015. Impacts of moso bamboo (Phyllostachys pubescens) invasion on dry matter and carbon and nitrogen stocks in a broad‐leaved secondary forest located in K yoto, western J apan. Plant Species Biol. 30, 81-95. https://doi.org/10.1111/1442-1984.12066\u003c/li\u003e\n\u003cli\u003eGallardo, B., Bacher, S., Barbosa, A.M., Gallien, L., Gonz\u0026aacute;lez-Moreno, P., Mart\u0026iacute;nez-Bolea, V., Sorte, C., Vimercati, G., Vil\u0026agrave;, M., 2024. Risks posed by invasive species to the provision of ecosystem services in Europe. Nat. Commun. 15, 2631. https://doi.org/10.1038/s41467-024-46818-3\u003c/li\u003e\n\u003cli\u003eGrayston, S.J., Wang, S., Campbell, C.D., Edwards, A.C., 1998. Selective influence of plant species on microbial diversity in the rhizosphere. Soil Biol. Biochem. 30, 369-378. https://doi.org/10.1016/S0038-0717(97)00124-7\u003c/li\u003e\n\u003cli\u003eGriffiths, B., Welschen, R.v., Van Arendonk, J., Lambers, H., 1992. The effect of nitrate-nitrogen supply on bacteria and bacterial-feeding fauna in the rhizosphere of different grass species. Oecologia 91, 253-259. https://doi.org/10.1007/BF00317793\u003c/li\u003e\n\u003cli\u003eHeděnec, P., Jim\u0026eacute;nez, J.J., Moradi, J., Domene, X., Hackenberger, D., Barot, S., Frossard, A., Oktaba, L., Filser, J., Kindlmann, P., 2022. Global distribution of soil fauna functional groups and their estimated litter consumption across biomes. Sci. Rep. 12, 17362. https://doi.org/10.1038/s41598-022-21563-z\u003c/li\u003e\n\u003cli\u003eHunt, H., Ingham, E., Coleman, D., Elliott, E., Reid, C., 1988. Nitrogen limitation of production and decomposition in prairie, mountain meadow, and pine forest. Ecology 69, 1009-1016. https://doi.org/10.2307/1941256\u003c/li\u003e\n\u003cli\u003eJones, T., Thompson, L., Lawton, J., Bezemer, T., Bardgett, R., Blackburn, T., Bruce, K., Cannon, P., Hall, G., Hartley, S., 1998. Impacts of rising atmospheric carbon dioxide on model terrestrial ecosystems. Science 280, 441-443. https://doi.org/10.1126/science.280.5362.441\u003c/li\u003e\n\u003cli\u003eKappes, H., Lay, R., Topp, W., 2007. Changes in different trophic levels of litter-dwelling macrofauna associated with giant knotweed invasion. Ecosystems 10, 734-744. https://doi.org/10.1007/s10021-007-9052-9\u003c/li\u003e\n\u003cli\u003eKassambara, A., 2019. Comparing groups: Numerical variables. Datanovia.[Google Scholar].\u003c/li\u003e\n\u003cli\u003eKhattak, W.A., Sun, J., Hameed, R., Zaman, F., Abbas, A., Khan, K.A., Elboughdiri, N., Akbar, R., He, F., Ullah, M.W., 2024. Unveiling the resistance of native weed communities: insights for managing invasive weed species in disturbed environments. Biol. Rev. 99, 753-777. https://doi.org/10.1111/brv.13043\u003c/li\u003e\n\u003cli\u003eLi, C., Bo, H., Song, B., Chen, X., Cao, Q., Yang, R., Ji, S., Wang, L., Liu, J., 2022. Reshaping of the soil microbiome by the expansion of invasive plants: shifts in structure, diversity, co-occurrence, niche breadth, and assembly processes. Plant Soil 477, 629-646. https://doi.org/10.1007/s11104-022-05445-x\u003c/li\u003e\n\u003cli\u003eLi, D., Wei, J., Wu, J., Zhong, Y., Chen, Z., He, J., Zhang, S., Yu, L., 2024. The expansion of moso bamboo (Phyllostachys edulis) forests into diverse types of forests in China from 2010 to 2020. Forests 15, 1418. https://doi.org/10.3390/f15081418\u003c/li\u003e\n\u003cli\u003eLi, Y., Ma, L., Wang, J., Zhang, J., 2021. Soil faunal community composition alters nitrogen distribution in different land use types in the Loess Plateau, China. Appl. Soil Ecol. 163, 103910. https://doi.org/10.1016/j.apsoil.2021.103910\u003c/li\u003e\n\u003cli\u003eLi, Y., Veen, G.C., Hol, W.G., Vandenbrande, S., Hannula, S.E., ten Hooven, F.C., Li, Q., Liang, W., Bezemer, T.M., 2020. \u0026lsquo;Home\u0026rsquo;and \u0026lsquo;away\u0026rsquo;litter decomposition depends on the size fractions of the soil biotic community. Soil Biol. Biochem. 144, 107783. https://doi.org/10.1016/j.soilbio.2020.107783\u003c/li\u003e\n\u003cli\u003eLiao, C., Peng, R., Luo, Y., Zhou, X., Wu, X., Fang, C., Chen, J., Li, B., 2008. Altered ecosystem carbon and nitrogen cycles by plant invasion: a meta‐analysis. New Phytol. 177, 706-714. https://doi.org/10.1111/j.1469-8137.2007.02290.x\u003c/li\u003e\n\u003cli\u003eLieurance, D., Cooper, A., Young, A.L., Gordon, D.R., Flory, S.L., 2018. Running bamboo species pose a greater invasion risk than clumping bamboo species in the continental United States. J. Nat. Conserv. 43, 39-45. https://doi.org/10.1016/j.jnc.2018.02.012\u003c/li\u003e\n\u003cli\u003eLin, D., Pang, M., Fanin, N., Wang, H., Qian, S., Zhao, L., Yang, Y., Mi, X., Ma, K., 2019. Fungi participate in driving home-field advantage of litter decomposition in a subtropical forest. Plant Soil 434, 467-480. https://doi.org/10.1007/s11104-018-3865-5\u003c/li\u003e\n\u003cli\u003eLiu, J., Compson, Z.G., Gui, X., Yang, Q., Song, Q., Huang, D., Ren, Z., Luan, F., 2024. Weak responses of soil microorganisms to leaf litter inputs after native Phyllostachys edulis invasion into adjacent native forests. Plant Soil 494, 685-699. https://doi.org/10.1007/s11104-023-06311-0\u003c/li\u003e\n\u003cli\u003eLiu, X., Siemann, E., Cui, C., Liu, Y., Guo, X., Zhang, L., 2019. Moso bamboo (Phyllostachys edulis) invasion effects on litter, soil and microbial PLFA characteristics depend on sites and invaded forests. Plant Soil 438, 85-99. https://doi.org/10.1007/s11104-019-04010-3\u003c/li\u003e\n\u003cli\u003eLong, K., Yin, R., Kardol, P., Wei, Q., Li, Y., Huang, J., 2023. Bamboo invasion alters Collembola community composition varying with life‐forms. Pest Manag. Sci. 79, 2517-2526. https://doi.org/10.1002/ps.7434\u003c/li\u003e\n\u003cli\u003eLuan, J., Liu, S., Li, S., Whalen, J.K., Wang, Y., Wang, J., Liu, Y., Dong, W., Chang, S.X., 2021. Functional diversity of decomposers modulates litter decomposition affected by plant invasion along a climate gradient. J. Ecol. 109, 1236-1249. https://doi.org/10.1111/1365-2745.13548\u003c/li\u003e\n\u003cli\u003eMa, A., Liu, H., Song, C., Tian, E., Wang, X., 2023. Home‐field advantage in litter decomposition: A critical review from a microbial perspective. J. Basic Microbiol. 63, 709-721. https://doi.org/10.1002/jobm.202200657\u003c/li\u003e\n\u003cli\u003eMacfadyen, A., 1953. Notes on methods for the extraction of small soil arthropods. J. Anim. Ecol., 65-77. https://doi.org/10.1016/S0031-4056(23)00341-4\u003c/li\u003e\n\u003cli\u003eMagoč, T., Salzberg, S.L., 2011. FLASH: fast length adjustment of short reads to improve genome assemblies. Bioinformatics 27, 2957-2963. https://doi.org/10.1093/bioinformatics/btr507\u003c/li\u003e\n\u003cli\u003eMelillo, J.M., Aber, J.D., Muratore, J.F., 1982. Nitrogen and lignin control of hardwood leaf litter decomposition dynamics. Ecology 63, 621-626. https://doi.org/10.2307/1936780\u003c/li\u003e\n\u003cli\u003eMilcu, A., Manning, P., 2011. All size classes of soil fauna and litter quality control the acceleration of litter decay in its home environment. Oikos 120, 1366-1370. https://doi.org/10.1111/j.1600-0706.2010.19418.x\u003c/li\u003e\n\u003cli\u003eMorris, S.J., Blackwood, C.B., 2024. The ecology of soil biota and their function, Soil microbiology, ecology and biochemistry. Elsevier, pp. 275-302. https://doi.org/10.1016/B978-0-12-822941-5.00010-7\u003c/li\u003e\n\u003cli\u003eNegrete-Yankelevich, S., Fragoso, C., Newton, A.C., Russell, G., Heal, O.W., 2008. Species-specific characteristics of trees can determine the litter macroinvertebrate community and decomposition process below their canopies. Plant Soil 307, 83-97. https://doi.org/10.1007/s11104-008-9585-5\u003c/li\u003e\n\u003cli\u003eOksanen, J., Blanchet, F., Kindt, R., Legendre, P., Minchin, P., O\u0026rsquo;hara, R., Simpson, G., Solymos, P., Stevens, M., Wagner, H., 2013a. Package \u0026lsquo;vegan\u0026rsquo;. Community Ecol. package, version 2.\u003c/li\u003e\n\u003cli\u003eOksanen, J., Blanchet, F.G., Kindt, R., Legendre, P., Minchin, P.R., O\u0026rsquo;hara, R., Simpson, G.L., Solymos, P., Stevens, M.H.H., Wagner, H., 2013b. Package \u0026lsquo;vegan\u0026rsquo;. Community Ecol. package, version 2, 1-295.\u003c/li\u003e\n\u003cli\u003eOlivia, C., Dumitru, M.T., 2023. THE FUNCTIONAL GROUPS OF INVERTEBRATES ACCORDING TO THEIR TROPHIC PREFERENCE. Oltenia, Stud. Comun. Stiintele Nat. 39.\u003c/li\u003e\n\u003cli\u003ePeng, Y., Holmstrup, M., Schmidt, I.K., De Schrijver, A., Schelfhout, S., Heděnec, P., Zheng, H., Bachega, L.R., Yue, K., Vesterdal, L., 2022. Litter quality, mycorrhizal association, and soil properties regulate effects of tree species on the soil fauna community. Geoderma 407, 115570. https://doi.org/10.1016/j.geoderma.2021.115570\u003c/li\u003e\n\u003cli\u003ePeng, Y., Schmidt, I.K., Zheng, H., Heděnec, P., Bachega, L.R., Yue, K., Wu, F., Vesterdal, L., 2020. Tree species effects on topsoil carbon stock and concentration are mediated by tree species type, mycorrhizal association, and N-fixing ability at the global scale. Forest Ecol. Manag. 478, 118510. https://doi.org/10.1016/j.foreco.2020.118510\u003c/li\u003e\n\u003cli\u003ePeng, Z., Lu, Y., Li, L., Zhao, Q., Feng, Q., Gao, Z., Lu, H., Hu, T., Yao, N., Liu, K., 2013. The draft genome of the fast-growing non-timber forest species moso bamboo (Phyllostachys heterocycla). Nat. Genet. 45, 456-461. https://doi.org/10.1038/ng.2569\u003c/li\u003e\n\u003cli\u003eP\u0026eacute;rez, G., Vila, M., Gallardo, B., 2022. Potential impact of four invasive alien plants on the provision of ecosystem services in Europe under present and future climatic scenarios. Ecosyst. Serv. 56, 101459. https://doi.org/10.1016/j.ecoser.2022.101459\u003c/li\u003e\n\u003cli\u003ePetersen, H., 1982. Basis for evaluation of the density and biomass data. Oikos 39, 295-305.\u003c/li\u003e\n\u003cli\u003ePhillips, R.P., Brzostek, E., Midgley, M.G., 2013. The mycorrhizal‐associated nutrient economy: a new framework for predicting carbon\u0026ndash;nutrient couplings in temperate forests. New Phytol. 199, 41-51. https://doi.org/10.1111/nph.12221\u003c/li\u003e\n\u003cli\u003ePremakumari, P.D., Sarayu, M.G., Das, G.G.M., Babu, K.V.D., Krishnan, R., Lawarence, B., Siraj, S.S., Pillai, A.R., Syamala, S.K.S., Murugan, K., 2022. Invasive exotic plant species and their influence on the environment, ecosystem services, economy and health: a search. J. Adv. Sci. Res. 13, 64-74. https://doi.org/10.55218/JASR.202213106\u003c/li\u003e\n\u003cli\u003eRaheem, A., Yohanna, P., Li, G., Noh, N.J., Iqbal, B., Tang, J., Du, D., Alahmadi, T.A., Ansari, M.J., Zhan, A., 2024. Unraveling the ecological threads: How invasive alien plants influence soil carbon dynamics. J. Environ. Manage. 356, 120556. https://doi.org/10.1016/j.jenvman.2024.120556\u003c/li\u003e\n\u003cli\u003eRai, A., Singh, A.K., Ghosal, N., Singh, N., 2016. Understanding the effectiveness of litter from tropical dry forests for the restoration of degraded lands. Ecol. Eng. 93, 76-81. https://doi.org/10.1016/j.ecoleng.2016.05.014 \u003c/li\u003e\n\u003cli\u003eRai, P.K., Singh, J., 2020. Invasive alien plant species: Their impact on environment, ecosystem services and human health. Ecol. Indic. 111, 106020. 10.1016/j.ecolind.2019.106020\u003c/li\u003e\n\u003cli\u003eReddy, H.R., Khan, M., Pasarkar, A., 2024. Soil Invertebrates As Sentinels Of Soil Health: A Zoological Approach To Soil Quality Assessment. Journal of Advanced Zoology 45. https://doi.org/10.53555/jaz.v45i1.3007\u003c/li\u003e\n\u003cli\u003eSauvadet, M., Chauvat, M., Brunet, N., Bertrand, I., 2017. Can changes in litter quality drive soil fauna structure and functions? Soil Biol. Biochem. 107, 94-103. https://doi.org/10.1016/j.soilbio.2016.12.018\u003c/li\u003e\n\u003cli\u003eSchaller, J., B\u0026ouml;ttger, R., Dudel, G., Ruess, L., 2016. Reed litter Si content affects microbial community structure and the lipid composition of an invertebrate shredder during aquatic decomposition. Limnologica 57, 14-22. https://doi.org/10.1016/j.limno.2015.12.002\u003c/li\u003e\n\u003cli\u003eScherber, C., Eisenhauer, N., Weisser, W.W., Schmid, B., Voigt, W., Fischer, M., Schulze, E.-D., Roscher, C., Weigelt, A., Allan, E., 2010. Bottom-up effects of plant diversity on multitrophic interactions in a biodiversity experiment. Nature 468, 553-556. https://doi.org/10.1038/nature09492\u003c/li\u003e\n\u003cli\u003eSeastedt, T., Crossley Jr, D., 1983. Nutrients in forest litter treated with naphthalene and simulated throughfall: a field microcosm study. Soil Biol. Biochem. 15, 159-165. https://doi.org/10.1016/0038-0717(83)90097-4\u003c/li\u003e\n\u003cli\u003eSet\u0026auml;l\u0026auml;, H., Marshall, V.G., Trofymow, J., 1996. Influence of body size of soil fauna on litter decomposition and 15N uptake by poplar in a pot trial. Soil Biol. Biochem. 28, 1661-1675. https://doi.org/10.1016/S0038-0717(96)00252-0 https://doi.org/10.1016/S0038-0717(96)00252-0\u003c/li\u003e\n\u003cli\u003eShannon, C.E., 2001. A mathematical theory of communication. ACM SIGMOBILE mobile computing and communications review 5, 3-55. https://doi.org/10.1145/584091.584093\u003c/li\u003e\n\u003cli\u003eShen, C., Gunina, A., Luo, Y., Wang, J., He, J.Z., Kuzyakov, Y., Hemp, A., Classen, A.T., Ge, Y., 2020. Contrasting patterns and drivers of soil bacterial and fungal diversity across a mountain gradient. Environ. Microbiol. 22, 3287-3301. https://doi.org/10.1111/1462-2920.15090\u003c/li\u003e\n\u003cli\u003eShokralla, S., Porter, T.M., Gibson, J.F., Dobosz, R., Janzen, D.H., Hallwachs, W., Golding, G.B., Hajibabaei, M., 2015. Massively parallel multiplex DNA sequencing for specimen identification using an Illumina MiSeq platform. Sci. Rep. 5, 1-7. https://doi.org/10.1038/srep09687\u003c/li\u003e\n\u003cli\u003eStrickland, M.S., Osburn, E., Lauber, C., Fierer, N., Bradford, M.A., 2009. Litter quality is in the eye of the beholder: initial decomposition rates as a function of inoculum characteristics. Funct. Ecol. 23, 627-636. https://doi.org/10.1111/j.1365-2435.2008.01515.x\u003c/li\u003e\n\u003cli\u003eSun, H., Hu, W., Dai, Y., Ai, L., Wu, M., Hu, J., Zuo, Z., Li, M., Yang, H., Ma, J., 2023. Moso bamboo (Phyllostachys edulis (Carri\u0026egrave;re) J. Houzeau) invasion affects soil microbial communities in adjacent planted forests in the Lijiang River basin, China. Front. Microbiol. 14, 1111498. https://doi.org/10.3389/fmicb.2023.1111498\u003c/li\u003e\n\u003cli\u003eTeam, R.C., 2013. R: A language and environment for statistical computing.\u003c/li\u003e\n\u003cli\u003eTeng, Q., Lu, X., Zhang, Q., Cai, L., Sardar, M.F., Li, Y., Abbas, T., Li, Y., Chang, S.X., Li, Y., 2023. Litterfall quality modulates soil ammonium and nitrate supply through altering microbial function in bamboo encroachment of broadleaf forests. Geoderma 437, 116592. https://doi.org/10.1016/j.geoderma.2023.116592\u003c/li\u003e\n\u003cli\u003eThukral, A.K., 2017. A review on measurement of Alpha diversity in biology. Agric. Res. J. 54. https://doi.org/10.5958/2395-146X.2017.00001.1\u003c/li\u003e\n\u003cli\u003eVerhoef, H., Brussaard, L., 1990. Decomposition and nitrogen mineralization in natural and agroecosystems: the contribution of soil animals. Biogeochemistry 11, 175-211. https://doi.org/10.1007/BF00004496\u003c/li\u003e\n\u003cli\u003eVil\u0026agrave;, M., Espinar, J.L., Hejda, M., Hulme, P.E., Jaro\u0026scaron;\u0026iacute;k, V., Maron, J.L., Pergl, J., Schaffner, U., Sun, Y., Py\u0026scaron;ek, P., 2011. Ecological impacts of invasive alien plants: a meta‐analysis of their effects on species, communities and ecosystems. Ecol. Lett. 14, 702-708. https://doi.org/10.1111/j.1461-0248.2011.01628.x\u003c/li\u003e\n\u003cli\u003eVivanco, L., Austin, A.T., 2008. Tree species identity alters forest litter decomposition through long‐term Plant Soil interactions in Patagonia, Argentina. J. Ecol. 96, 727-736. https://doi.org/10.1111/j.1365-2745.2008.01393.x\u003c/li\u003e\n\u003cli\u003eWang, J., Wang, Y., He, N., Ye, Z., Chen, C., Zang, R., Feng, Y., Lu, Q., Li, J., 2020. Plant functional traits regulate soil bacterial diversity across temperate deserts. Sci. Total Environ. 715, 136976. https://doi.org/10.1016/j.scitotenv.2020.136976\u003c/li\u003e\n\u003cli\u003eWardle, D., 2002. Communities and ecosystems: linking the aboveground and belowground components. https://doi.org/10.1515/9781400847297\u003c/li\u003e\n\u003cli\u003eWei, Q., Rui, Y., Huang, J., Vogler, A.P., Li, Y., Miao, X., Kardolg, P., 2021. The diversity of soil mesofauna decline after bamboo invasion in subtropical China. Sci. Total Environ., 147982. https://doi.org/10.1016/j.scitotenv.2021.147982\u003c/li\u003e\n\u003cli\u003eWeidlich, E.W., Fl\u0026oacute;rido, F.G., Sorrini, T.B., Brancalion, P.H., 2020. Controlling invasive plant species in ecological restoration: A global review. J. Appl. Ecol. 57, 1806-1817. https://doi.org/10.1111/1365-2664.13656\u003c/li\u003e\n\u003cli\u003eWickham, H., 2011. ggplot2. Wiley interdisciplinary reviews: computational statistics 3, 180-185. http://dx.doi.org/10.1002/wics.1637\u003c/li\u003e\n\u003cli\u003eWolfe, B.E., Klironomos, J.N., 2005. Breaking new ground: soil communities and exotic plant invasion. BioScience 55, 477-487. https://doi.org/10.1641/0006-3568(2005)055[0477:BNGSCA]2.0.CO;2\u003c/li\u003e\n\u003cli\u003eWRB, 2006. A framework for international classification, correlation and communication. World Soil Resour. Rep. 103.\u003c/li\u003e\n\u003cli\u003eXiao, H., Liu, J., Li, F., 2023. Both alpha and beta diversity of nematode declines in response to moso bamboo expansion in south China. Appl. Soil Ecol. 183, 104761. https://doi.org/10.1016/j.apsoil.2022.104761\u003c/li\u003e\n\u003cli\u003eXu, Q.-F., Liang, C.-F., Chen, J.-H., Li, Y.-C., Qin, H., Fuhrmann, J.J., 2020. Rapid bamboo invasion (expansion) and its effects on biodiversity and soil processes+. Glob. Ecol. Conserv. 21, e00787. https://doi.org/10.1016/j.gecco.2019.e00787\u003c/li\u003e\n\u003cli\u003eYang, K., Zhu, J., Zhang, W., Zhang, Q., Lu, D., Zhang, Y., Zheng, X., Xu, S., Wang, G.G., 2022. Litter decomposition and nutrient release from monospecific and mixed litters: Comparisons of litter quality, fauna and decomposition site effects. J. Ecol. 110, 1673-1686. https://doi.org/10.1111/1365-2745.13902\u003c/li\u003e\n\u003cli\u003eYang, X., Chen, J., 2009. Plant litter quality influences the contribution of soil fauna to litter decomposition in humid tropical forests, southwestern China. Soil Biol. Biochem. 41, 910-918. https://doi.org/10.1016/j.soilbio.2008.12.028\u003c/li\u003e\n\u003cli\u003eYang, Y., Wu, Q., Yang, W., Wu, F., Zhang, L., Xu, Z., Liu, Y., Tan, B., Li, H., Zhou, W., 2020. Temperature and soil nutrients drive the spatial distributions of soil macroinvertebrates on the eastern Tibetan Plateau. Ecosphere 11, e03075. https://doi.org/10.1002/ecs2.3075\u003c/li\u003e\n\u003cli\u003eYin, R., Gruss, I., Eisenhauer, N., Kardol, P., Thakur, M.P., Schmidt, A., Xu, Z., Siebert, J., Zhang, C., Wu, G.-L., 2019. Land use modulates the effects of climate change on density but not community composition of Collembola. Soil Biol. Biochem. 138, 107598. https://doi.org/10.1016/j.soilbio.2019.107598\u003c/li\u003e\n\u003cli\u003eYin, X., Song, B., Dong, W., Xin, W., Wang, Y., 2010. A review on the eco-geography of soil fauna in China. J. Geogr. Sci. 20, 333-346. https://doi.org/10.1007/s11442-010-0333-4\u003c/li\u003e\n\u003cli\u003eYu, D.W., Ji, Y., Emerson, B.C., Wang, X., Ye, C., Yang, C., Ding, Z., 2012. Biodiversity soup: metabarcoding of arthropods for rapid biodiversity assessment and biomonitoring. Methods Ecol. Evol. 3, 613-623. https://doi.org/10.1111/j.2041-210X.2012.00198.x\u003c/li\u003e\n\u003cli\u003eZhang, L., 2023a. Bamboo Expansion and Soil Microbial Communities, Bamboo Expansion: Processes, Impacts, and Management. Springer, pp. 197-208.\u003c/li\u003e\n\u003cli\u003eZhang, L., 2023b. Bamboo Expansion Into Adjacent Ecosystems, Bamboo Expansion: Processes, Impacts, and Management. Springer, pp. 19-37.\u003c/li\u003e\n\u003cli\u003eZhao, C., Li, Y., Zhang, C., Miao, Y., Liu, M., Zhuang, W., Shao, Y., Zhang, W., Fu, S., 2021. Considerable impacts of litter inputs on soil nematode community composition in a young Acacia crassicapa plantation. Soil Ecol. Lett. 3, 145-155. https://doi.org/10.1007/s42832-021-0085-3\u003c/li\u003e\n\u003cli\u003eZhong, Y., Yan, W., Wang, R., Wang, W., Shangguan, Z., 2018. Decreased occurrence of carbon cycle functions in microbial communities along with long-term secondary succession. Soil Biol. Biochem. 123, 207-217. https://doi.org/10.1016/j.soilbio.2018.05.017\u003c/li\u003e\n\u003cli\u003eZhou, S., Butenschoen, O., Barantal, S., Handa, I.T., Makkonen, M., Vos, V., Aerts, R., Berg, M.P., McKie, B., Van Ruijven, J., 2020. Decomposition of leaf litter mixtures across biomes: The role of litter identity, diversity and soil fauna. J. Ecol. 108, 2283-2297. https://doi.org/10.1111/1365-2745.13452\u003c/li\u003e\n\u003cli\u003eZhu, M., Fanin, N., Wang, Q., Xu, Z., Liang, S., Ye, J., Lin, F., Yuan, Z., Mao, Z., Wang, X., 2024. High functional breadth of microbial communities decreases home-field advantage of litter decomposition. Soil Biol. Biochem. 188, 109232. https://doi.org/10.1016/j.soilbio.2023.109232\u003c/li\u003e\n\u003c/ol\u003e"},{"header":"Table","content":"\u003cp\u003eTable 2 is available in the Supplementary Files section.\u003c/p\u003e\n"}],"fulltextSource":"","fullText":"","funders":[],"hasAdminPriorityOnWorkflow":false,"hasManuscriptDocX":true,"hasOptedInToPreprint":true,"hasPassedJournalQc":"","hasAnyPriority":false,"hideJournal":false,"highlight":"","institution":"","isAcceptedByJournal":true,"isAuthorSuppliedPdf":false,"isDeskRejected":"","isHiddenFromSearch":false,"isInQc":false,"isInWorkflow":true,"isPdf":false,"isPdfUpToDate":true,"isWithdrawnOrRetracted":false,"journal":{"display":true,"email":"[email protected]","identity":"plant-and-soil","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":false,"externalIdentity":"plso","sideBox":"Learn more about [Plant and Soil](https://www.springer.com/journal/11104)","snPcode":"11104","submissionUrl":"https://submission.nature.com/new-submission/11104/3","title":"Plant and Soil","twitterHandle":"","acdcEnabled":true,"dfaEnabled":true,"editorialSystem":"em","reportingPortfolio":"Springer Hybrid","inReviewEnabled":true,"inReviewRevisionsEnabled":false},"keywords":"Plant invasion, Litter reciprocal transplants, Soil fauna communities, Litter quality, Feeding guilds","lastPublishedDoi":"10.21203/rs.3.rs-6625409/v1","lastPublishedDoiUrl":"https://doi.org/10.21203/rs.3.rs-6625409/v1","license":{"name":"CC BY 4.0","url":"https://creativecommons.org/licenses/by/4.0/"},"manuscriptAbstract":"\u003ch2\u003eBackground and aims\u003c/h2\u003e \u003cp\u003eInvasive plant species represent significant threats to ecosystems globally, disrupting native habitats and adversely affecting biodiversity and ecosystem functions and services. This study focused on the impact of bamboo invasion, specifically the spread of moso bamboo (\u003cem\u003ePhyllostachys edulis\u003c/em\u003e), on soil fauna communities in subtropical forests.\u003c/p\u003e\u003ch2\u003eMethods\u003c/h2\u003e \u003cp\u003eUsing litter reciprocal transplant experiments between uninvaded broadleaf forests and invaded bamboo forests, we investigated how alterations in litter quality influenced soil fauna diversity and community composition.\u003c/p\u003e\u003ch2\u003eResults\u003c/h2\u003e \u003cp\u003eReciprocal litter transplants primarily affected soil fauna community composition through species replacement without changing their diversity or ecological function. Notably, bamboo litter, which is considered high quality, had a more pronounced impact on soil fauna communities in low-quality litter environments but not vice versa.\u003c/p\u003e\u003ch2\u003eConclusion\u003c/h2\u003e \u003cp\u003eThese findings highlight the intricate relationship between litter quality, soil fauna communities, and ecosystem stability in the context of plant invasion. Understanding these dynamics provides valuable insights for ecosystem management strategies in the face of invasive plant species.\u003c/p\u003e","manuscriptTitle":"Litter quality drives the shift of soil fauna structure without changing its ecological function","msid":"","msnumber":"","nonDraftVersions":[{"code":1,"date":"2025-06-09 06:48:35","doi":"10.21203/rs.3.rs-6625409/v1","editorialEvents":[{"type":"communityComments","content":0},{"type":"decision","content":"Major revisions","date":"2025-07-13T08:28:08+00:00","index":"","fulltext":""},{"type":"reviewerAgreed","content":"","date":"2025-06-05T22:18:53+00:00","index":0,"fulltext":""},{"type":"reviewersInvited","content":"","date":"2025-06-02T22:38:06+00:00","index":"","fulltext":""},{"type":"editorInvited","content":"Plant and Soil","date":"2025-05-09T23:59:23+00:00","index":"","fulltext":""},{"type":"editorAssigned","content":"","date":"2025-05-09T07:40:04+00:00","index":"","fulltext":""},{"type":"submitted","content":"Plant and Soil","date":"2025-05-09T02:04:20+00:00","index":"","fulltext":""}],"status":"published","journal":{"display":true,"email":"[email protected]","identity":"plant-and-soil","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":false,"externalIdentity":"plso","sideBox":"Learn more about [Plant and Soil](https://www.springer.com/journal/11104)","snPcode":"11104","submissionUrl":"https://submission.nature.com/new-submission/11104/3","title":"Plant and Soil","twitterHandle":"","acdcEnabled":true,"dfaEnabled":true,"editorialSystem":"em","reportingPortfolio":"Springer Hybrid","inReviewEnabled":true,"inReviewRevisionsEnabled":false}}],"origin":"","ownerIdentity":"9f481760-d328-42e2-becf-be0075acc3ab","owner":[],"postedDate":"June 9th, 2025","published":true,"recentEditorialEvents":[],"rejectedJournal":[],"revision":"","amendment":"","status":"published-in-journal","subjectAreas":[],"tags":[],"updatedAt":"2025-09-08T16:05:06+00:00","versionOfRecord":{"articleIdentity":"rs-6625409","link":"https://doi.org/10.1007/s11104-025-07828-2","journal":{"identity":"plant-and-soil","isVorOnly":false,"title":"Plant and Soil"},"publishedOn":"2025-09-03 15:57:31","publishedOnDateReadable":"September 3rd, 2025"},"versionCreatedAt":"2025-06-09 06:48:35","video":"","vorDoi":"10.1007/s11104-025-07828-2","vorDoiUrl":"https://doi.org/10.1007/s11104-025-07828-2","workflowStages":[]},"version":"v1","identity":"rs-6625409","journalConfig":"researchsquare"},"__N_SSP":true},"page":"/article/[identity]/[[...version]]","query":{"redirect":"/article/rs-6625409","identity":"rs-6625409","version":["v1"]},"buildId":"8U1c8b4HqxoKbykW_rLl7","isFallback":false,"isExperimentalCompile":false,"dynamicIds":[84888],"gssp":true,"scriptLoader":[]}