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Article

Analysis of Understory Plant Community Assembly Differences in Moso Bamboo Forests in the Subtropical Evergreen Broad-Leaved Forest Region of Eastern China

by
Zhiwei Ge
1,2,
Tao Yu
2,
Xuying Tian
2,
Xiangxiang Chen
2,
Yiwen Yao
2 and
Lingfeng Mao
1,2,*
1
Co-Innovation Center for Sustainable Forestry in Southern China, Nanjing 210037, China
2
College of Ecology and Environment, Nanjing Forestry University, Nanjing 210037, China
*
Author to whom correspondence should be addressed.
Forests 2025, 16(3), 478; https://doi.org/10.3390/f16030478
Submission received: 4 February 2025 / Revised: 27 February 2025 / Accepted: 7 March 2025 / Published: 8 March 2025
(This article belongs to the Special Issue Sustainable Management of Forest Stands)
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Abstract

:
Moso bamboo (Phyllostachys edulis (Carrière) J. Houz.) forests are a vital forest type in subtropical China. This study investigates the diversity, floristic composition, and phylogenetic structure of understory vegetation in these bamboo forests within evergreen broad-leaved forests of eastern subtropical China. Using grid-based sampling, we calculated species diversity and phylogenetic indices, and employed correlation analysis, redundancy analysis, and structural equation modeling to assess the effects of canopy closure, soil properties, and topography. The understory exhibited high species richness, with shrub layer demonstrating phytogeographic characteristics predominantly associated with tropical distribution types, while the herbaceous layer is characterized by temperate distribution types. Canopy closure and environmental factors significantly influenced shrub diversity, showing a clustered phylogenetic structure (NTI > 0, NRI > 0) and a negative correlation with species diversity. In contrast, the herb layer displayed a divergent phylogenetic structure (NTI < 0, NRI < 0), shaped by neutral stochastic processes, reflecting endemic taxa and interspecific interactions. These findings emphasize the need for targeted management practices to conserve understory biodiversity, focusing on enhancing floristic and phylogenetic diversity while protecting endemic species and their ecological interactions.

1. Introduction

Forests cover approximately one-third of the global terrestrial area and serve as critical reservoirs of biodiversity [1]. China’s subtropical regions, accounting for 25% of the nation’s land area, constitute a globally significant biodiversity hotspot characterized by abundant endemic species and representative subtropical evergreen broad-leaved forest ecosystems [2,3]. These ecosystems play a vital role in maintaining endemic biota conservation and ecosystem services in East Asia [4]. As a transitional zone between northern and central subtropical climatic regimes, the region’s complex topography and pronounced climatic gradients provide valuable insights into species distribution patterns, community organization, and biogeographical processes [5]. The eastern subtropical region has become a biodiversity hotspot facing intense anthropogenic pressures from rapid socioeconomic development [6]. However, evolving environmental policies have facilitated the transformation of abandoned plantations into semi-natural ecosystems. With Moso bamboo maintaining dominance amid shifting management priorities [7], its ecological role has evolved from economic production to multifaceted ecosystem service provision [8,9]. While previous studies traditionally consider Moso bamboo plantations as suboptimal for biodiversity conservation [10], emerging evidence suggests that natural succession over 20–30 years post abandonment, combined with low-intensity management, could challenge this perspective [11,12]. These dynamics necessitate a comprehensive reassessment of Moso bamboo forests as transitional ecosystems in anthropogenic landscapes.
Biodiversity fundamentally sustains ecosystem services in plantations [13], where understory vegetation serves as a key component in maintaining ecological structure and function [14]. Current research on Moso bamboo forests predominantly examines limited spatial scales, lacking comprehensive analysis of understory floristic distribution and genetic diversity. The ecological significance of these forests largely stems from their understory vegetation, which sustains forest structure and ecosystem resilience while enhancing energy flow and nutrient cycling [15,16,17]. Notably, studies suggest that subtropical plantations with minimal disturbance can harbor substantial biological resources [18,19,20], highlighting the urgency for large-scale investigations into understory vegetation composition in Moso bamboo forests. Modern diversity assessment methods now incorporate phylogenetic analyses, enabling evaluation of both species relationships and ecosystem characteristics [21,22,23]. As evolution is an irreversible and unpredictable process, conserving phylogenetic diversity is crucial for preserving the potential for future biodiversity evolution. This approach provides a scientific foundation for understanding biodiversity distribution patterns and formulating effective conservation strategies [24,25,26]. Therefore, applying phylogenetic analysis to investigate species composition and community assembly mechanisms in Moso bamboo forests provides critical insights for understanding their dynamic ecological value and advancing sustainable forest management strategies under changing environmental conditions.
Moso bamboo spans over half of China’s provinces, with Zhejiang and Fujian containing the largest distributions [9]. These low-elevation hilly regions serve as traditional forest resource hubs and ecological barriers in eastern subtropical China [27]. While the biodiversity conservation potential of these areas requires thorough investigation [28,29], research on understory plant diversity in local Moso bamboo forests remains limited despite documented ecological impacts from natural forest conversion to plantations [30,31,32]. Although studies confirm Moso bamboo’s influence on understory vegetation composition [33], its effects on community assembly mechanisms across plant life forms remain unclear.
Building on current ecological evidence, we hypothesize that shrub communities in Moso bamboo forests are predominantly structured by niche-based assembly processes, where environmental filtering drives the coexistence of species with analogous ecological niches through trait convergence [34,35]. This mechanistic dominance reflects shrubs’ superior adaptive capacity to canopy-mediated constraints. In contrast, herbaceous communities likely adhere to neutral stochastic assembly processes, characterized by weak environmental filtering and probabilistic colonization dynamics under bamboo canopy regimes [36]. Therefore, we focus on representative Moso bamboo forests in the Zhejiang and Fujian provinces, situated within the evergreen broad-leaved forest regions of eastern subtropical China. By analyzing species/phylogenetic diversity patterns and environmental drivers across plant life forms, we aim to reveal community assembly mechanisms. Our findings evaluate these plantations’ conservation potential and inform sustainable management strategies for anthropogenic forest ecosystems.

2. Materials and Methods

2.1. Study Site and Design

This study was conducted across the Zhejiang and Fujian provinces, situated within the evergreen broad-leaved forest regions of eastern subtropical China. A uniform grid method was employed to partition the study area into 100 km × 100 km units, with field surveys conducted in selected grids. Sampling plots and survey points were established as outlined in the Figure 1 and Table A1 (Appendix A). Field investigations were carried out from June to September 2021, covering site selection, route planning, and plot establishment. In each county, representative Moso bamboo forest communities were selected, encompassing diverse topographical features, habitats, and elevation gradients. A total of thirty 20 m × 20 m tree layer plots were established. Within each tree layer plot, five-point sampling was used to set up 2 m × 2 m shrub layer plots and 1 m × 1 m herb layer plots, resulting in 150 plots for each layer. Survey data included measurements of individual trees with a diameter at breast height (DBH) ≥ 3 cm, recording DBH, tree height, and canopy width. Shrub layer data encompassed coverage, abundance, basal diameter, and average height. For herbaceous plants, coverage, average height, and the number of individuals (or clumps) were documented. Species names, abundance, and richness were recorded, and specimens were collected for identification. Community characteristics and environmental factors of the larger plots were also documented.
Topographical factors (TF) included altitude (Alt), canopy density (Cd), aspect (Asp), slope position (PS), and slope degree (Slo), derived from field surveys. Soil physicochemical factors (SF) included pH, bulk density (BD), cation exchange capacity (CEC), total nitrogen (TN), total phosphorus (TP), total potassium (TK), and soil organic carbon (SOC). Soil data were obtained from the National Earth System Science Data Center, accessible via the National Science and Technology Resource Sharing Service Platform (http://www.geodata.cn, accessed on 6 June 2024).

2.2. Study Method

2.2.1. Geographical Distribution Analysis of Vascular Plant Families and Genera

Floristic analysis of families and genera of ferns and seed plants was conducted based on the works of Zang [39], Wu et al. [40], Li et al. [41], the Flora of China website (https://www.iplant.cn/, accessed on 13 December 2024), and the Plants of the World Online database (https://powo.science.kew.org/, accessed on 13 December 2024).

2.2.2. Calculation of Phylogenetic Diversity Indices

Building upon the hypotheses formulated in this study, phylogenetic analytical approaches—by virtue of their taxon-independent nature in reflecting evolutionary diversity dimensions—provide a scientifically robust framework for informing conservation area prioritization and strategy optimization [42,43]. The specific phylogenetic metrics employed in this investigation, along with their computational algorithms, are systematically detailed as follows.
First, the plantlist package in R 4.0.3 [44] was used to validate species names in each plot. Species names were standardized based on the CTPL Plant Species Checklist System, with corrections or removals applied to account for changes in scientific names, as well as variations or subspecies. This process yielded a database of 387 plant species for the study. Subsequently, the V.Phylomaker package [45] was employed to construct a phylogenetic tree for Moso bamboo forests (see Figure S1 in Supplementary Materials). Using the curated species database and the global mega-phylogeny (GBOTB.extended.tre) proposed by Smith et al. [46]. As the backbone, the root and basal node information for the largest clusters at the genus or family level were extracted. The phylo.maker function in scenario 3 (scenarios = “S3”) was used to generate a comprehensive phylogenetic tree for the bamboo forests, which was then utilized to calculate phylogenetic diversity indices.
Phylogenetic diversity (PD) at the community level was quantified using Faith’s [47] metric, which measures the total branch length of the phylogenetic tree. The net relatedness index (NRI) and nearest taxon index (NTI) were used to assess community phylogenetic structure. Using the constructed phylogenetic tree, the picante package [48] was applied to compute phylogenetic distances for all species in each plot (ses.MPD and ses.MNTD). A null model provided by the picante package was used to randomize species names in the species pool 999 times, generating distributions of ses.MPD and ses.MNTD under the null model. Ses.MPD equals the negative of NRI, and ses.MNTD equals the negative of NTI. When NRI and NTI are greater than 0, it indicates phylogenetic clustering compared to the null model. When NRI and NTI are less than 0, it indicates phylogenetic overdispersion (divergence) compared to the null model. When NRI and NTI are equal to 0, it indicates a random phylogenetic structure compared to the null model.
Generally, NRI is considered more sensitive to the overall phylogenetic clustering or evenness, while NTI is more sensitive to the clustering and evenness near the tips of the phylogenetic tree [49]. NRI describes the overall phylogenetic structure of species in the community, whereas NTI focuses on the interspecific phylogenetic relationships of the nearest neighbors [50]. The formulas for calculating NTI and NRI are as follows [51,52]:
N T I = − 1 × M N T D o b s e r v e d − M N T D r a n d o m i z e d s d M N T D r a n d o m i z e d
N R I = − 1 × M P D o b s e r v e d − M P D r a n d o m i z e d s d M P D r a n d o m i z e d
where MNTDobserved and MPDobserved are the observed mean nearest taxon distance and mean pairwise distance between species in the community, respectively, and MNTDrandomized and MPDrandomized are the expected mean values of MNTD and MPD after 999 randomizations under the null model. The terms sdMNTDrandomized and sdMPDrandomized represent the standard deviations of MNTD and MPD, respectively, after randomization.

2.2.3. Measurement of Species Diversity

In this study, species richness (SR), the Shannon diversity index (H), the Simpson diversity index (D), and the Pielou evenness index (J) were used to measure species diversity [53].

2.2.4. Statistical Analysis

Diversity indices and redundancy analysis were calculated in R software (version 4.0.3) using the vegan package [54]. Pearson’s correlation analysis between understory diversity indices and environmental factors was performed using the psych package [55]. A structural equation model (SEM) was constructed using least squares estimation via the plspm package [56] to analyze the effects of biotic and abiotic factors on community diversity indices. All data were normalized prior to analysis, and figures were generated using the ggplot2 package [57].

3. Results

3.1. Basic Characteristics of Moso Bamboo Forests and Geographical Floristic Analysis of Understory Vegetation

3.1.1. Basic Characteristics of Moso Bamboo Forests

This study surveyed a total land area of 229,500 km2 (including 19,300 km2 of bamboo forests) in the subtropical Zhejiang and Fujian provinces, eastern China, with detailed sampling across 12,000 m2 revealing 387 understory plant species from 107 families and 232 genera within the bamboo ecosystems (Table 1). The tree layer comprised 23 families, 29 genera, and 33 species. Excluding the dominant species, Phyllostachys edulis, the Lauraceae family exhibited the highest species richness (three species), followed by the genera Diospyros, Litsea, Castanopsis, and Heptapleurum, each represented by two species. In the shrub layer, 73 families, 148 genera, and 268 species were identified. The Fabaceae family had the highest species count (20 species), followed by Rosaceae (18 species) and both Fagaceae and Lauraceae (16 species each). At the genus level, Rubus was the most species-rich genus (12 species), followed by Ilex, Quercus, and Ficus, each with four species. The herb layer included 47 families, 85 genera, and 110 species. Poaceae was the most species-rich family (13 species), followed by Asteraceae (eight species) and both Polygonaceae and Urticaceae (six species each). At the genus level, Persicaria had the highest number of species (five species), followed by Selaginella and Dioscorea, each with four species.

3.1.2. Distribution Types of Dominant Genera in Understory Vegetation of Moso Bamboo Forests

Based on the classification of plant genus distribution types, the vascular plants in the shrub layer of the bamboo forests in the eastern subtropical evergreen broad-leaved forest region of China comprised 148 genera. Among these, 10 genera containing five or more species were classified as dominant genera (Table 2), totaling 64 species and accounting for approximately 24% of the shrub layer species. Tropical distribution genera were the most abundant, including six genera: Ilex, Ficus, Callicarpa, Symplocos, Eurya, and Machilus. Temperate distribution genera included three genera: Quercus, Castanopsis, and Actinidia. One genus with a worldwide distribution, Rubus, was also recorded. The distribution types of shrub layer plants were relatively diverse, with tropical genera dominating.
For herbaceous layer vascular plants, 85 genera were recorded. The six genera containing three or more species were classified as dominant genera (Table 3), including three temperate distribution genera: Persicaria, Dioscorea, and Polygonatum; two genera with worldwide distributions: Selaginella and Carex; and one tropical distribution genus, Cyclosorus. The distribution types in the herbaceous layer were more complex, with temperate and widespread distribution types dominating, while tropical genera were less prevalent.
Unique genera are defined as those containing only one species, and unique species are those appearing only once in the sampled plots. In the shrub layer, there were 91 unique genera, accounting for 61.49% of the total genera, and 143 unique species, accounting for 53.36% of the total species. In the herbaceous layer, there were 70 unique genera, representing 82.35% of the total genera, and 63 unique species, representing 57.27% of the total species. The proportions of unique species and genera were higher in the herbaceous layer than in the shrub layer (Table 4).

3.2. Species and Phylogenetic Diversity Characteristics of Understory Vegetation in Moso Bamboo Forests

Species diversity is a direct measure of richness and biological diversity within a community (Table 5). The species richness and Shannon diversity index of understory shrub-layer plants were higher than those of the herbaceous layer, whereas the Simpson index was lower in the shrub layer compared to the herbaceous layer. Interspecific relationships within the community may lead to another form of diversity difference, known as phylogenetic diversity, which reflects the potential evolutionary and community assembly relationships among species to some extent [58]. Studies have shown that, in the shrub layer, NTI > 0 and NRI > 0, indicating that community assembly is primarily driven by environmental filtering. In contrast, the phylogenetic structure indices for the herbaceous layer showed NRI < 0, with NTI exhibiting both positive and negative trends, suggesting that NRI is more sensitive to the overall phylogenetic structure’s aggregation and uniformity within the community [50,59]. Thus, the phylogenetic structure of the shrub layer is more aggregated, driven mainly by environmental filtering, whereas the herbaceous layer’s phylogenetic structure is more divergent, driven primarily by interspecific competition.

3.3. Environmental Influences on Understory Plant Diversity

3.3.1. Correlation and Redundancy Analysis of Environmental Factors and Understory Plant Diversity

Correlation analysis between various factors and diversity indices (Figure 2) revealed that, in the understory shrub layer, altitude significantly and negatively affected species richness (SR) (p < 0.05). Soil total nitrogen (TN) (p < 0.001) and organic carbon (SOC) (p < 0.05) significantly and negatively influenced the Shannon index (p < 0.001). Soil total phosphorus (TP), TN, total potassium (TK), pH, cation exchange capacity (CEC), bulk density (BD), canopy closure (Cd), and slope all significantly and positively affected the Simpson index. TN, TK, SOC, pH, and BD significantly and negatively influenced the Pielou index. The mean nearest taxon phylogenetic distance (MNTD) was significantly and positively correlated with SOC. In the herbaceous layer, only altitude significantly affected the Simpson index and MNTD, showing a negative correlation with the Simpson index and a positive correlation with MNTD.
Redundancy analysis (RDA) was performed to explore the effects of environmental factors on diversity patterns in the shrub and herbaceous layers (Figure 3). In the shrub layer, RDA1 and RDA2 explained 35.76% and 30.76% of the variation, respectively. The analysis indicated that soil factors significantly influenced the diversity pattern of understory shrub vegetation in bamboo forests, with TN (R2 = 0.522), CEC (R2 = 0.470), and SOC (R2 = 0.336) having highly significant effects (p < 0.01), while BD (R2 = 0.678), pH (R2 = 0.590), and TK (R2 = 0.283) also significantly influenced the diversity gradient (p < 0.05). Among the topographical factors, slope and canopy closure (Cd) also exhibited significant effects. Additionally, the Shannon index (R2 = 0.942), SR (R2 = 0.847), phylogenetic diversity (PD) (R2 = 0.730), and Pielou index (R2 = 0.687) were highly correlated with environmental factors (p < 0.001), indicating that these diversity indices effectively explain the variation in community diversity. In the herbaceous layer, RDA1 and RDA2 explained 55.87% and 25.74% of the variation, respectively. Only altitude showed marginal significance with some diversity indices, such as MNTD (R2 = 0.191, p = 0.06). Diversity indices significantly explained the variation in community diversity (p < 0.05), with SR, Shannon, and Simpson indices demonstrating strong consistency in explaining the variation. Based on the correlation analysis results, the diversity of shrub layer plants is strongly influenced by canopy closure, topographical factors, and soil environmental factors, whereas herbaceous layer plants are primarily associated with altitude.

3.3.2. Structural Equation Modeling of Environmental Factors and Understory Plant Diversity

A partial least squares (PLS) method was used to construct structural equation models (SEMs) to further analyze the influence of environmental factors on understory plant diversity. After bootstrap correction, the models exhibited a good fit (GoF A = 0.459, GoF B = 0.439) (Figure 4A,B). For the shrub layer, canopy closure (Cd) showed a highly significant positive effect on soil factors (S.F.) (path coefficient, PC = 0.689, p < 0.001) and species diversity (Div.) (PC = 0.637, p < 0.001), while S.F. (PC = −0.599) and topographical factors (T.F.) (PC = −0.431) exhibited significant negative effects on Div. (p < 0.001). Div. emerged as a key negative predictor of phylogenetic diversity (Phylodiv.) (PC = −0.93, p < 0.001). Path decomposition revealed direct effects of Cd (PC = 0.64), S.F. (PC = −0.60), and T.F. (PC = −0.43) on Div., with indirect effects from Cd (PC = −0.41) and T.F. (PC = −0.01). For Phylodiv., Div. had a strong direct negative effect (PC = −0.93), while Cd (direct: PC = 0.10; indirect: PC = −0.17), S.F. (direct: PC = 0.05; indirect: PC = 0.55), and T.F. (direct: PC = 0.06; indirect: PC = 0.41) showed mixed influences (Figure 4C,E).
In the herbaceous layer, Cd and T.F. were key drivers of the soil environment (p < 0.001), but environmental factors showed no significant direct effects on diversity indices. Div. negatively influenced Phylodiv. (PC = −0.72, p < 0.001), while S.F. marginally affected Div. (PC = 0.58, p = 0.078). Path decomposition indicated that Cd (direct: PC = −0.14; indirect: PC = 0.23) and T.F. (direct: PC = −0.06; indirect: PC = 0.39) influenced Div. through indirect pathways, and Phylodiv. was primarily driven by Div. (PC = −0.72), with additional minor effects from Cd (direct: PC = 0.10; indirect: PC = 0.01), S.F. (direct: PC = 0.20; indirect: PC = −0.42), and T.F. (direct: PC = −0.37; indirect: PC = −0.10) (Figure 4D,F).
Overall, shrub layer diversity was directly enhanced by canopy closure (PC = 0.637) but suppressed by soil (PC = −0.599) and topographical factors (PC = −0.431), whereas herbaceous layer diversity remained largely unaffected by environmental factors. Critically, species diversity negatively predicted phylogenetic diversity in both layers (PC = −0.93 for shrubs; PC = −0.72 for herbs), highlighting a trade-off between taxonomic and evolutionary diversity.

4. Discussion

4.1. Understory Plant Diversity and Floristic Characteristics in Moso Bamboo Forests

This study investigates the species diversity of Moso bamboo forests within subtropical evergreen broad-leaved forests of eastern China, documenting 107 families, 232 genera, and 387 plant species. Among these, 106 families, 226 genera, and 377 species were recorded in the understory vegetation, representing over 90% of families and approximately 97% of genera and species. Understory vegetation constitutes a critical component of the plant resource pool in Moso bamboo forests, playing an essential roles in ecological functions, species renewal, and forest ecosystem regulation, particularly in artificial forest settings [60]. These findings highlight the importance of understory vegetation in biodiversity conservation and underscore its value as a reservoir of genetic resources [61].
The results demonstrate that the understory flora in these bamboo forests retains significant diversity, preserving substantial plant genetic and phylogenetic resources. Compared to typical subtropical evergreen and deciduous forests in eastern China, which host over 150 families, 520 genera, and 1600 vascular plant species [62], our study sites retained 70% of family-level diversity and 23% of species-level diversity, demonstrating notable retention capacity despite bamboo dominance. Dominant families (Leguminosae, Rosaceae, Fagaceae) contrasted with 70% of families represented by less than three species, revealing a heterogeneous community structure [63].
For species within the same genus, the genus as a taxonomic unit typically evolves and diverges from a common ancestor, forming a monophyletic group. In evolutionary biology, species within a genus exhibit relatively stable and unified traits yet maintain distinct internal variations. In biogeography and taxonomy, the genus level is considered appropriate for classification [40,41]. Therefore, analyzing the distribution types and floristic composition of genera is of significant importance for regional floristic research. Taxonomic analysis revealed evolutionary and biogeographic significance, with genus-level distributions aligning with subtropical transitional characteristics. Shrub layers predominantly contained tropical-affiliated genera, while herbaceous layers featured temperate-affiliated genera—consistent with the “Out of the Tropics” hypothesis of tropical origins for temperate biodiversity [64,65]. The observed floristic composition suggests a strong capacity to preserve diversity in this region [66], challenging the notion of bamboo forests as “green deserts” and instead emphasizing their role in sustaining plant diversity under appropriate management [67].
The conservation paradox of Moso bamboo forests requires urgent resolution, given their dominance in a global biodiversity hotspot. These transitional ecosystems demonstrate dual dynamics: serving as secondary habitats for native species recruitment while risking biotic homogenization through bamboo expansion. This ecological ambiguity necessitates longitudinal studies to determine whether abandoned plantations ultimately enhance or compromise regional biodiversity, particularly given increasing global plantation abandonment [12].
Our findings mandate targeted strategies prioritizing understory vegetation conservation as genetic reservoirs, coupled with continuous monitoring of biodiversity hotspots and unique lineages. While providing critical baseline data, this study underscores the need for long-term spatiotemporal monitoring to distinguish persistent successional trends from transient states, combined with multidimensional analyses to unravel community dynamics in heterogeneous landscapes.

4.2. Environmental Drivers of Understory Plant Diversity and Differences in Community Assembly Mechanisms

Canopy closure regulates understory vegetation development and composition by modulating light availability, temperature, humidity, and water distribution. This study demonstrates that canopy closure significantly increased the Simpson index of the shrub layer (p < 0.001) but had no measurable effect on herbaceous layer diversity. Increased canopy closure reduces understory gap frequency, altering microclimatic conditions and selectively filtering plant species adapted to low-light environments [68]. These dynamics likely explain the observed phylogenetic clustering in the shrub layer (NTI > 0, NRI > 0) versus the divergence in the herbaceous layer (NRI < 0), suggesting canopy closure acts as a primary environmental filter structuring shrub communities.
Soil physicochemical properties critically influence nutrient availability and species diversity. In the shrub layer, SOC positively correlated with MNTD, indicating that soil-mediated environmental filtering shapes species assembly. TN and CEC further significantly reduced shrub diversity indices (p < 0.01), consistently with findings that soil carbon enrichment promotes understory cover but reduces species richness [60,69]. Conversely, herbaceous layer diversity showed only marginal sensitivity to soil factors (p = 0.078), likely due to bamboo root systems altering surface soil properties [70] and competitive exclusion limiting herbaceous niche space [71,72].
Topographic factors (slope, aspect, altitude) indirectly shape diversity through light and water redistribution [73]. Slope positively influenced the shrub layer’s Simpson index but negatively affected species richness, while altitude weakly modulated herbaceous layer’s metrics. These patterns align with studies showing stronger topographic effects on shrub communities due to their deeper root systems and broader environmental tolerances compared to herbs [74,75,76]. Aspect-driven microclimate variations further amplify compositional differences across vegetation types.
Phylogenetic diversity analyses revealed contrasting assembly mechanisms between layers. The shrub layer’s clustered structure (NTI > 0, NRI > 0) reflects environmental filtering under niche conservatism, where canopy and soil factors restrict species to phylogenetically related lineages with shared adaptations [77,78]. In contrast, the herbaceous layer’s divergent structure (NRI < 0) suggests competition-driven exclusion of close relatives, consistent with neutral processes dominating early successional stages [59,79,80]. This diversity–phylogeny decoupling aligns with global patterns of stochastic herbaceous assembly [81,82].
Endemism patterns further highlight community assembly contrasts. The herbaceous layer’s higher endemic genera (82.35%) and species (57.27%) versus the shrub layer (61.49%, 53.36%) supports neutral theory predictions of stochastic dominance in herb communities [83,84]. Early successional herbaceous species may exploit spatial priority effects, perpetuating random colonization and inhibiting succession. In contrast, the shrub layer’s environmental filtering aligns with niche theory, where abiotic factors hierarchically structure communities [85].
These findings underscore the dual role of Moso bamboo forests as both ecological filters and biodiversity reservoirs. Targeted management balancing canopy density and soil conservation could enhance shrub layer diversity, while herbaceous communities may require disturbance regimes to maintain stochastic assembly processes. Future studies should quantify long-term successional trajectories and anthropogenic impacts on these dynamics. This also confirms the hypothesis proposed in this study that there are indeed differences in the community assembly processes between shrub layer plants and herbaceous layer plants in the understory vegetation of the Moso bamboo forest ecosystem.

4.3. Biodiversity Conservation Potential of Moso Bamboo Forests

This study reveals that subtropical Moso bamboo forests exhibit a notable capacity to conserve understory plant diversity, preserving both temperate and tropical floristic elements alongside widespread species. These forests maintain rich phytogeographic diversity, challenging the prevailing narrative that focuses predominantly on the negative impacts of Moso bamboo invasion on biodiversity [86]. Emerging evidence suggests that alternative ecological roles of bamboo ecosystems warrant attention. For instance, Liu et al. [67] demonstrate that managed bamboo forests under enclosure policies can enhance biodiversity patterns, while Chen et al. [87] found comparable shrub and herb layer richness between bamboo forests and native ecosystems. Ma et al. [88] further emphasized integrating regional biogeographic history into conservation planning to protect latent diversity. These insights underscore the need for expanded research on bamboo forests’ biodiversity conservation potential, particularly regarding phylogenetic diversity of ecologically significant species. Incorporating evolutionary history into conservation strategies [89]—where phylogenetic diversity reflects evolutionary distinctiveness independent of taxonomic status—could strengthen the scientific basis for reserve design and management optimization [42,43].
A pivotal finding is the contrasting phylogenetic assembly mechanisms between shrub and herb layers. The shrub layer’s clustered structure (NTI > 0, NRI > 0) reflects environmental filtering, whereas the herb layer’s divergent structure (NRI < 0) arises from competitive exclusion and stochastic colonization. Phylogenetically divergent communities, though less explored, may enhance biodiversity conservation by harboring distantly related lineages with unique functional traits [26]. While artificial forests are often dismissed as biodiversity-poor, this study highlights their underestimated phylogenetic conservation value—a dimension frequently overlooked in conventional analyses focusing on climate [90], understory structure [91], or soil properties [92]. Echoing Norden et al. [93], we emphasize that long-term forest diversity restoration requires attention to intrinsic community dynamics. Our results demonstrate that understory community assembly operates semi-independently from overstory controls: shrub communities are structured by abiotic filters, while herbaceous communities exhibit neutral assembly patterns characterized by spatial priority effects and competitive preemption [36]. These divergent pathways create heterogeneous evolutionary trajectories within the understory, suggesting that sub-canopy vegetation may harbor greater diversity than traditionally assumed [94].
This mechanistic understanding demands transformative management approaches for Moso bamboo forests. Moving beyond viewing them merely as invasive threats, acknowledging their dual capacity as ecological filters and biodiversity reservoirs enables more nuanced conservation strategies. Specifically, practices that balance canopy density control with soil preservation could enhance shrub layer diversity, while maintaining herb layer stochasticity through regulated disturbances may conserve their distinct assembly patterns. Future research must prioritize long-term monitoring of successional trajectories and anthropogenic impacts, particularly on evolutionarily vulnerable lineages. Paradoxically, intensifying anthropogenic pressures in subtropical hotspots—driven by rapid socioeconomic development—have catalyzed novel ecological pathways as plantations transition toward semi-natural ecosystems. This widespread landscape transformation urgently requires scientific examination of evolving ecosystem services and their mechanisms in human-mediated successional systems.

5. Conclusions

Moso bamboo forests in China’s eastern subtropical evergreen broadleaf forests maintain a relatively diverse understory plant community. The shrub layer demonstrates tropical affinity in dominant genera composition, contrasting with the temperate-origin species dominating the herbaceous layer, resulting in multifaceted phytogeographic characteristics. Canopy closure, soil properties, and topographic features were identified as primary determinants of shrub-layer species diversity. Phylogenetic analyses demonstrated significant clustering in the shrub layer (NTI > 0; NRI > 0), confirming environmental filtering as the predominant assembly mechanism, where elevated species diversity corresponded with reduced phylogenetic diversity. Conversely, herbaceous communities exhibited limited environmental correlations with divergent phylogenetic patterns (NTI < 0; NRI < 0), indicative of stochastic processes and biotic interactions governing community organization. These results necessitate refined conservation strategies prioritizing understory vegetation in Moso bamboo ecosystems, particularly through structured monitoring programs to protect habitat-specialized taxa.
These insights should inform adaptive management protocols for subtropical biodiversity hotspots, where understanding ecosystem service trajectories in management-transformed plantations requires balanced integration of ecological conservation and utilization imperatives. Furthermore, the established framework serves as a critical reference for global regions undergoing analogous silvicultural transitions, particularly in anthropogenically modified landscapes demanding scientifically grounded strategies to achieve economically viable and ecologically sustainable forest management practices.

Supplementary Materials

The following supporting information can be downloaded at: https://www.mdpi.com/article/10.3390/f16030478/s1, Figure S1: The phylogenetic tree of the overall species pool surveyed in this study.

Author Contributions

Conceptualization, Z.G. and T.Y.; methodology, T.Y., Y.Y., and X.T.; validation, T.Y. and X.T.; formal analysis, T.Y.; investigation, Y.Y. and X.C.; writing—original draft preparation, T.Y.; writing—review and editing, Z.G., T.Y., and L.M.; visualization, T.Y.; supervision, L.M. and Z.G. All authors have read and agreed to the published version of the manuscript.

Funding

This work was funded by the “Jiangsu Forestry Science & Technology Innovation and Extension Project (Project No: LYKJ 2022 01)”, the “National Key Research and Development Projects of the Ministry of Science and Technology of China (2023YFF0805800)”, the “Jiangsu Social Development Project (BE2022792)”, and “The National Natural Science Foundation of China (grant numbers 31870506, 32271712)”.

Data Availability Statement

The datasets presented in this article are not readily available because the data are part of other ongoing studies.

Acknowledgments

The authors wish to thank the members of the Biodiversity and Ecological Conservation Research Group at Nanjing Forestry University for their assistance with sampling and data analysis in this study. The data support from the “National Earth System Science Data Center, National Science & Technology Infrastructure of China (https://www.geodata.cn)” is also acknowledged. The authors would also like to thank reviewers and the editors for their valuable input in relation to this article.

Conflicts of Interest

The authors declare no conflicts of interest.

Appendix A

Table A1. Characteristics of the plots.
Table A1. Characteristics of the plots.
ProvincePlot NumberDominate Tree SpeciesAltitude/mCrown
Density		AspectLongitude and Latitude
ZhejiangMZ01Phyllostachys edulis137.50.6Southwest slopeE:120°34′10″ N:28°48′37″
MZ022100.9Northeast slopeE:120°35′20″ N:28°37′32″
MZ031800.85Northeast slopeE:120°35′12″ N:28°38′23″
MZ041540.65West slopeE:119°27′10″ N:30°14′25″
MZ054480.7East slopeE:119°26′27″ N:30°19′29″
MZ063150.65East slopeE:119°26′32″ N:30°18′43″
MZ073300.7Southeast slopeE:118°37′02″ N:28°43′56″
MZ083560.65South slopeE:118°36′58″ N:28°43′51″
MZ093780.7Northeast slopeE:118°36′47″ N:28°43′48″
MZ102540.75Southeast slopeE:119°39′47″ N:27°55′31″
MZ112300.7South slopeE:119°39′55″ N:27°55′28″
MZ122100.6Southwest slopeE:119°39′18″ N:27°55′16″
MZ136280.67Northwest slopeE:118°39′22″ N:28°29′33″
MZ147500.75Southeast slopeE:118°42′26″ N:28°30′59″
MZ151840.8North slopE:118°33′37″ N:28°32′37″
FujianMZ162910.6West slopeE:117°15′27″ N:24°30′56″
MZ173860.85North slopeE:117°16′08″ N:24°30′34″
MZ184330.7Southwest slopeE:117°14′43″ N:24°31′27″
MZ19931.20.65East slopeE:117°31′13″ N:25°57′06″
MZ201172.70.7Northeast slopeE:117°29′54″ N:25°57′51″
MZ217100.65Southeast slopeE:117°33′14″ N:25°57′19″
MZ222800.6Northwest slopeE:117°50′23″ N:27°38′56″
MZ23178.60.6North slopeE:117°47′53″ N:27°37′19″
MZ24278.50.7East slopeE:117°48′55″ N:27°38′17″
MZ256620.75Northeast slopeE:118°46′47″ N:26°22′43″
MZ267600.68North slopeE:118°49′34″ N:26°22′25″
MZ276300.7Northwest slopeE:118°46′39″ N:26°22′44″
MZ284460.6Southeast slopeE:117°51′47″ N:26°49′37″
MZ292190.6Southwest slopeE:117°56′26″ N:26°51′24″
MZ30231.50.6South slopeE:117°46′06″ N:26°45′09″

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Forests 16 00478 g001
Figure 1. Geographic location map and sampling sites in the study. Note: the distribution of bamboo forests shown in the figure is primarily derived from Li et al. [37] and Chen [38]. The map review number for this figure is GS(2024)0650 from the National Platform for Common GeoSpatial Information Services, PRC.
Figure 1. Geographic location map and sampling sites in the study. Note: the distribution of bamboo forests shown in the figure is primarily derived from Li et al. [37] and Chen [38]. The map review number for this figure is GS(2024)0650 from the National Platform for Common GeoSpatial Information Services, PRC.
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Forests 16 00478 g002
Figure 2. Correlation between environmental factors and diversity indices. (a) Shrub layer; (b) herbaceous layer. Asterisks indicate significance levels: *: p < 0.05, **: p < 0.01, and ***: p < 0.001.
Figure 2. Correlation between environmental factors and diversity indices. (a) Shrub layer; (b) herbaceous layer. Asterisks indicate significance levels: *: p < 0.05, **: p < 0.01, and ***: p < 0.001.
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Forests 16 00478 g003
Figure 3. Redundancy analysis of environmental factors and diversity indices. (a) Shrub layer; (b) herbaceous layer.
Figure 3. Redundancy analysis of environmental factors and diversity indices. (a) Shrub layer; (b) herbaceous layer.
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Forests 16 00478 g004
Figure 4. Structural equation model of environmental factors and diversity index in understory shrub layer (A,C,E) and herbaceous layer (B,D,F). Note: S.F., soil factors, T.F., topographical factors, Div., species diversity indices, Phylodiv., phylogenetic diversity indices. Asterisks indicate significance levels: ***: p < 0.001.
Figure 4. Structural equation model of environmental factors and diversity index in understory shrub layer (A,C,E) and herbaceous layer (B,D,F). Note: S.F., soil factors, T.F., topographical factors, Div., species diversity indices, Phylodiv., phylogenetic diversity indices. Asterisks indicate significance levels: ***: p < 0.001.
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Table 1. Overview of understory plants and key species importance values in Moso bamboo forests.
Table 1. Overview of understory plants and key species importance values in Moso bamboo forests.
Shrub LayerHerbaceous Layer
Species
Profile		SpeciesImportant ValueSpecies
Profile		SpeciesImportant Value
73 families,
148 genera,
268 species		Maesa japonica2.8747 families,
85 genera,
110 species		Woodwardia japonica10.16
Rubus buergeri2.84Dicranopteris pedata9.00
Camellia sinensis2.27Lophatherum gracile8.97
Pternandra caerulescens2.24Selaginella moellendorffii4.47
Loropetalum chinense2.22Dryopteris fuscipes2.83
Table 2. Dominant genera of vascular plants in the shrub layer of Moso bamboo forests in the eastern subtropical evergreen broadleaf forest region of China.
Table 2. Dominant genera of vascular plants in the shrub layer of Moso bamboo forests in the eastern subtropical evergreen broadleaf forest region of China.
Serial NumberGenusNumber of
Species in the
Region		Number of
Species in China		Percentage of China’s Total SpeciesNumber of
Species in the World		Percentage of World’s Total SpeciesPercentage of the Region’s Total SpeciesDistribution Type
1Rubus121946.197001.713.17Worldwide distribution
2Ilex72043.434201.671.85Discontinuous distribution between tropical Asia and tropical America
3Quercus75113.733002.331.85Discontinuous distribution between East Asia and North America
4Ficus7987.1410000.701.85Pantropical distribution
5Callicarpa64613.041903.161.59Pantropical distribution
6Castanopsis5637.941204.171.32Discontinuous distribution between East Asia and North America
7Actinidia5529.62559.091.32East Asia (eastern Himalayas—Japan)
8Symplocos5776.493001.671.32Discontinuous distribution between tropical Asia, Oceania (to New Zealand), and Central to South America (or Mexico)
9Eurya5816.171303.851.32Discontinuous distribution between tropical Asia and tropical America
10Machilus5687.351005.001.32West Malaysia, Central Malaysia, East Malaysia
Table 3. Dominant genera of vascular plants in the herb layer of Moso bamboo forests in the eastern subtropical evergreen broadleaf forest region of China.
Table 3. Dominant genera of vascular plants in the herb layer of Moso bamboo forests in the eastern subtropical evergreen broadleaf forest region of China.
Serial NumberGenusNumber of
Species in the Region		Number of
Species in China		Percentage of China’s Total SpeciesNumber of
Species in the World		Percentage of World’s Total SpeciesPercentage of the Region’s Total SpeciesDistribution Type
1Persicaria51134.422302.171.32Discontinuous distribution between East Asia and North America
2Selaginella4725.567000.571.06Worldwide distribution
3Dioscorea4498.166000.671.06East Asia (eastern Himalayas—Japan)
4Cyclosorus31272.362501.200.79Pantropical distribution
5Carex35270.5720000.150.79Worldwide distribution
6Polygonatum3319.68407.500.79Northern temperate distribution
Table 4. Overview of endemic genera and species of different life forms in the understory of Moso bamboo forests in the eastern subtropical evergreen broadleaf forest region of China.
Table 4. Overview of endemic genera and species of different life forms in the understory of Moso bamboo forests in the eastern subtropical evergreen broadleaf forest region of China.
Frequency of Occurrence
(%)		Frequency of Occurrence
(count)		Shrub LayerHerbaceous Layer
NumberPercentage (%)NumberPercentage (%)
Genera3.33≤19161.497082.35
6.67≤212383.117992.94
Species3.33≤114353.366357.27
6.67≤212546.648072.73
Table 5. Plant species and phylogenetic diversity in the understory of Moso bamboo forests in the eastern subtropical evergreen broadleaf forest region of China.
Table 5. Plant species and phylogenetic diversity in the understory of Moso bamboo forests in the eastern subtropical evergreen broadleaf forest region of China.
IndexShrub LayerHerbaceous Layer
Species diversity indexSR22.93 ± 1.149.93 ± 0.77
H2.68 ± 0.071.77 ± 0.07
D0.89 ± 0.010.76 ± 0.02
J0.87 ± 0.020.80 ± 0.02
Phylogenetic diversity indexPD2441.66 ± 95.991773.95 ± 88.84
MPD258.97 ± 6.06533.53 ± 13.92
MNTD150.07 ± 4.55240.34 ± 14.21
NTI1.32 ± 0.15−0.07 ± 0.20
NRI1.25 ± 0.11−2.63 ± 0.17
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Ge, Z.; Yu, T.; Tian, X.; Chen, X.; Yao, Y.; Mao, L. Analysis of Understory Plant Community Assembly Differences in Moso Bamboo Forests in the Subtropical Evergreen Broad-Leaved Forest Region of Eastern China. Forests 2025, 16, 478. https://doi.org/10.3390/f16030478

AMA Style

Ge Z, Yu T, Tian X, Chen X, Yao Y, Mao L. Analysis of Understory Plant Community Assembly Differences in Moso Bamboo Forests in the Subtropical Evergreen Broad-Leaved Forest Region of Eastern China. Forests. 2025; 16(3):478. https://doi.org/10.3390/f16030478

Chicago/Turabian Style

Ge, Zhiwei, Tao Yu, Xuying Tian, Xiangxiang Chen, Yiwen Yao, and Lingfeng Mao. 2025. "Analysis of Understory Plant Community Assembly Differences in Moso Bamboo Forests in the Subtropical Evergreen Broad-Leaved Forest Region of Eastern China" Forests 16, no. 3: 478. https://doi.org/10.3390/f16030478

APA Style

Ge, Z., Yu, T., Tian, X., Chen, X., Yao, Y., & Mao, L. (2025). Analysis of Understory Plant Community Assembly Differences in Moso Bamboo Forests in the Subtropical Evergreen Broad-Leaved Forest Region of Eastern China. Forests, 16(3), 478. https://doi.org/10.3390/f16030478

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