Spatial Patterns of Michelia macclurei and Mytilaria laosensis plantations after 24 Years of Near-Natural Transformation

Spatial Patterns of Michelia macclurei and Mytilaria laosensis plantations after 24 Years of Near-Natural Transformation | 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 Spatial Patterns of Michelia macclurei and Mytilaria laosensis plantations after 24 Years of Near-Natural Transformation Ziyi Chen, Liuqi Feng, Yifan Liu, Yuanfa Li This is a preprint; it has not been peer reviewed by a journal. https://doi.org/ 10.21203/rs.3.rs-8701430/v1 This work is licensed under a CC BY 4.0 License Status: Under Review Version 1 posted 4 You are reading this latest preprint version Abstract The spatial pattern of plantations changes after near-natural transformation, but the underlying driving mechanisms remain unclear. To address this issue, we established three permanent plots in monoculture and mixed plantations of native tree species Mytilaria laosensis Lecomte and Michelia macclurei Dandy in southern China. Trees were classified into planted trees and regenerations, and regenerations were further divided into shade-tolerant and shade-intolerant functional groups based on light adaptability. We applied the univariate pair correlation function g 11 ( r ) to analyze the distribution patterns of planted trees and regenerations, and bivariate pair correlation function g 12 ( r ) to analyze spatial correlations between planted trees and regenerations, as well as between functional groups. Results indicate: (1) planted trees exhibited aggregated distributions in all stands at small spatial scales ( r = 0–2 m, r = 0–3 m), regenerations showed aggregation across a wider range of spatial scales; (2) spatial correlations between planted trees and regenerations were positive in the Mytilaria laosensis Lecomte monoculture stand and the mixed stand ( r = 0–2 m), while it was negatively correlated in monoculture stand of Michelia macclurei Dandy ( r = 2–20 m); (3) spatial correlations between shade-tolerant and shade-intolerant regenerations was negatively correlated in all stands, but at slightly different scales ( r = 0–2 m, r = 0–3 m, r = 0–5 m). These results suggest that planting patterns and functional traits jointly drive spatial pattern shifts in plantations, leading to increased spatial complexity within stands and deepening our understanding of the near-natural transformation process of plantations. Plantation Planting pattern Distribution pattern Spatial correlation Functional groups Figures Figure 1 Figure 2 Figure 3 Figure 4 Figure 5 1. Introduction Spatial patterns serve as a crucial method for investigating the relationship between community structure and ecological processes, with their core encompassing the spatial distribution of individuals and interspecies correlations (Getzin et al., 2008 ). Distribution patterns reveal the degree of aggregation within communities across multiple scales through the distribution of tree points, reflecting the effects of seed dispersal limitations, heterogeneous resource availability, and historical disturbances and succession (Pommerening, 2002 ). Spatial correlations describe the spatial relationships among individuals from different populations, life stages, or origins, providing important evidence for interpreting the maintenance of species diversity, niche differentiation, and community stability (Getzin et al., 2008 ; Ben-Said, 2021 ). Spatial patterns have been extensively used to elucidate the mechanisms of species diversity formation and coexistence in natural forests (Stoll and Newbery, 2005 ; Gu et al., 2019 ), but rarely used to analyze the succession and regeneration of spatial structure in plantations (Ben-Said, 2021 ). However, with the advancement of ecological civilization construction and the deepening of the concept of multifunctional forest utilization, the management of monoculture forests guided by maximizing timber production and economic benefits have gradually exposed problems such as insufficient ecosystem service functions (Paquette and Messier, 2010 ; Liu et al., 2018 ). Many stands are undergoing near-natural transformation in order to adjust stand structure and enhance ecosystem functions (Ming et al., 2020 ; O’Hara, 2016 ), yet the transformation process and underlying drivers remain unclear. Applying spatial pattern analysis to plantations research helps deepen our understanding of community structural dynamics and provides scientific support for optimizing management strategies (Forrester, 2014 ; Zhang et al., 2019 ). Planting patterns, namely monoculture stands and mixed stand, influence the spatial patterns of plantations under natural succession. The species composition, spatial arrangement, and proportions of planted trees directly determine the intensity of neighborhood interactions (Forrester, 2014 ; Pretzsch and Schütze, 2021 ). Individual plants in monoculture forests have similar resource requirements, and the intense competition within their neighborhoods makes them prone to self-thinning (Hui et al., 2018 ), which alters the spatial patterns. In contrast, differences in growth processes and resource utilization among species in mixed stand may alleviate or alter interaction ways between neighboring species (Forrester, 2014 ; Pretzsch and Schütze, 2021 ). Meanwhile, planting patterns affect seed composition and sources (Cole et al., 2010 ; Liu et al., 2022 ), thereby altering the establishment processes and spatial distribution patterns of regenerations (Huanca Nuñez et al., 2021 ; Wang et al., 2019 ). In monoculture stands, parent trees are dominated by a single species, and regenerations mainly originates from seed rain of planted trees and soil seed banks (Seidler and Plotkin, 2006 ; Liu et al., 2022 ). By contrast, parent trees in mixed stand are more diverse, expanding seed sources and dispersal range, better regulating the forest microclimate, increasing the quantity and quality of litter (Huang et al., 2017 ; Zhang et al., 2022 ), and promoting the regeneration of site-quality-sensitive species (De Frenne et al., 2019 ). Furthermore, due to complex structure, mixed stand shows self-regulation and resistance to disturbance during succession (Liu et al., 2018 ; Pretzsch and Zenner, 2017 ), which increases the differences in spatial structure between mixed and monoculture stands. In addition to planting patterns, the functional traits of regeneration also influence the spatial patterns of plantations (Asanok et al., 2013 ; Rahayu et al., 2022 ). Specifically, differences among functional groups in resource acquisition and utilization affect regeneration processes and further modify spatial patterns (Forrester, 2014 ; Zambrano et al., 2020 ). Although many factors influence the renewal process, light may be the most important limiting factor at small spatial scales (De Lombaerde et al., 2020 ; Sangsupan et al., 2021 ). Shade-tolerant groups have lower light compensation points and higher light-use efficiency (Valladares and Niinemets, 2008 ), which enables them to maintain photosynthesis and growth under low understory light. Therefore, they readily form small-scale aggregations beneath or near the canopy of planted trees (Getzin et al., 2008 ; Paluch, 2024 ). In contrast, shade-intolerant groups are highly dependent on light, prefer canopy gaps or forest edges, and their spatial patterns are closer to dispersed or random distributions (Klopčič et al., 2015 ). Current research on plantations mainly examines regeneration dynamics from the perspectives of species composition and environmental drivers, focusing on the effects of habitat heterogeneity and competitive pressure (Ali et al., 2019 ; Kremer et al., 2022 ). However, few studies analyze the role of regeneration functional groups in shifts in stand spatial patterns. The subtropical region of southern China boasts favorable hydrothermal conditions and numerous native tree species occur, which play important roles in regional ecological restoration, timber production, and landscape construction (Lu et al., 2017 ; Li et al., 2021c ). In some areas, monoculture and mixed stands of native species have been established for decades, but most stands were harvested once the planted trees reach economic maturity. Using Mytilaria laosensis Lecomte and Michelia macclurei Dandy stands that have exceeded one rotation period (20 a) as examples, this study examines how planting patterns influence the spatial patterns of native-species plantations and hypothesizes: (1) Mortality of planted trees and the occurrence of regeneration (Zhang et al., 2019 ; Li et al., 2021c ) shift the stand point pattern from an initially regular distribution to a random or aggregated distribution. Moreover, seed dispersal modes and dispersal limitation affect the distribution of regeneration (Nathan and Muller-Landau, 2000 ; Dent and Estrada-Villegas, 2021 ), leading to aggregation (Hypothesis 1); (2) Regeneration establishment depends strongly on the overstory vegetation and the microenvironment it creates (O’Brien et al., 2007 ; Rautio et al., 2023 ), therefore planted trees and regenerations attract each other spatially (Hypothesis 2); (3) Differences in the plant morphology and vertical structure of planted trees between monoculture and mixed stands (Montgomery and Chazdon, 2001 ; Pretzsch, 2014 ), cause functional groups with different light dependence to occupy different spatial positions (Ligot et al., 2014 ; Forrester et al., 2018 ), resulting in negative correlations between functional groups (Hypothesis 3). To test these hypotheses, we established permanent plots in a Mytilaria laosensis monoculture stand (ML stand), a Michelia macclurei monoculture stand (MM-Da stand), and M. laosensis – M. macclurei mixed stand (Mixed stand) in southern Guangxi, and used census data for validation. The results will elucidate the drivers of change in spatial patterns and the regeneration mechanisms of functional groups, thereby promoting near-natural transformation of plantations. 2. Materials and Methods 2.1 Regional Overview The experimental site is located at the Jiepai substation of Gaofeng Forest Farm, Guangxi, southern China (108°21′–108°24′ E, 22°56′–24°00′ N). The region has a warm climate with abundant rainfall and sufficient sunshine, and it is characterized as a southern subtropical monsoon climate. The average annual temperature is 21.56°C, the annual precipitation is approximately 2056 mm, and the average annual sunshine duration is 1551 h. The landform is dominated by low mountains and hills with gentle relief. The parent rock is mainly granite, and long-term weathering has resulted in lateritic red soils and purple soils, with an average soil depth exceeding 80 cm and good water and fertilizer retention properties. Native tree species in the study area primarily form the overstory, including Mytilaria laosensis Lecomte, Machilus macclurei Dandy, and Castanopsis hystrix Hook. f. and Thomson ex A. DC., among others. In addition, the understory species composition is diverse, common shrubs include Melicope pteleifolia (Champ. ex Benth.) Hartley, Ardisia quinquegona Blume, Wendlandia uvariifolia Hance, Actinodaphne pilosa (Lour.) Merr., Desmos chinensis Lour., Lonicera confusa (Sweet) DC., and Psychotria asiatica L., while common herbaceous species include Sarcandra glabra (Thunb.) Nakai, Sarcophyton taiwanianum (Hayata) Garay, Miscanthus floridulus (Lab.) Warb. ex Schum. et Laut., and Dicranopteris dichotoma (Thunb.) Bernh, among others (Yang et al., 2025 ). 2.2 Plots establishment and Data collection The specific experimental site was located on a gentle slope with an inclination of approximately 25°. In 1999, we clear-cut a mature Chinese fir ( Cunninghamia lanceolata (Lamb.) Hook.) plantation and then established monoculture stands of Mytilaria laosensis , Michelia macclurei , and their mixed stand, with an approximate spacing of 3.5 m × 3.5 m. The stem ratio of Mytilaria laosensis to Michelia macclurei in the mixed stand was approximately 2:1. Manual tending was conducted during the first 1–5 years after planting, after which the stands were in a state of natural succession and gradually developed a multispecies coexistence pattern (Yang et al., 2025 ). In 2024–2025, we established three permanent plots with sizes of 140 m × 40 m, 70 m × 50 m, and 70 m × 40 m, according to the shape and area of each class (Fig. 1 ). First, we used a total station (model NTS-372R 10 , South Surveying and Mapping Company, Guangzhou, China) to define plot boundaries and divided each plot into n subplots of 10 m × 10 m. Then, we mapped the coordinates ( x , y , z ) of all trees within each plot (diameter at breast height, DBH ≥ 1.0 cm; DBH < 1.0 cm and height ≥ 0.5 m). Finally, each tree was tagged to identify the species and to measure its dimensions. The measured variables included DBH (m), tree height (m), crown width (m), and ground diameter (cm) for non-planted species with DBH ≥ 1.0 cm or DBH < 1.0 cm and height ≥ 0.5 m, these individuals included almost all planted trees and regenerations in the plots (Yang et al., 2025 ). 2.3 Data Analysis 2.3.1 Classification of Tree Groups To better analyze spatial patterns in monoculture and mixed plantations and the regeneration characteristics of functional groups, we classified trees into planted trees and regenerations based on their origin, and further divided regenerations into shade-tolerant and shade-intolerant functional groups based on their light adaptability. When counting plants, sprouted individuals of planted trees below a height of 1.3 m were counted as independent plants. In ML Stand, we recorded a total of 889 individuals, including 488 planted trees, 219 shade-tolerant regenerations, and 182 shade-intolerant regenerations. In MM-Da Stand, we recorded 1318 individuals in total, including 208 planted trees, 500 shade-tolerant regenerations, and 610 shade-intolerant regenerations. In Mixed Stand, we recorded 1397 individuals, including 201 planted trees, 667 shade-tolerant regenerations, and 529 shade-intolerant regenerations (Table 1 ). Table 1 Basic information on tree populations in ML stand, MM-Da stand, ML and MM-Da mixed stand. Others refer to populations with N < 10. N, number of individuals; DBH, diameter at breast height; TH, tree height; CW, crown width. Plot Type Species Number DBH ± sd(cm) TH ± sd(m) CW ± sd(m) ML stand planted trees ML 488 22.39 ± 6.7 20.36 ± 4.8 3.95 ± 2.05 reg.(shade-tolerant) AP 26 0.80 ± 0.44 0.91 ± 0.55 0.25 ± 0.18 ML 145 3.00 ± 3.12 4.08 ± 3.44 0.89 ± 0.86 MM-Da 23 1.04 ± 0.78 1.13 ± 0.77 0.29 ± 0.3 Others 25 1.22 ± 0.69 1.26 ± 0.76 0.31 ± 0.25 reg.(shade-intolerant) AP 25 0.58 ± 0.20 0.86 ± 0.39 0.2 ± 0.09 ML 98 2.47 ± 2.63 3.90 ± 2.79 0.83 ± 0.75 MM-Da 32 1.10 ± 0.82 1.05 ± 0.71 0.3 ± 0.26 Others 27 1.12 ± 0.85 1.25 ± 0.99 0.4 ± 0.38 MM-Da stand planted trees MM-Da HH 206 2 21.42 ± 7.7 36.91 ± 24.8 19.92 ± 4.3 16.0 ± 5.6 4.29 ± 1.96 reg.(shade-tolerant) AP 42 1.31 ± 0.88 1.60 ± 1.13 0.37 ± 0.21 AQ 54 0.99 ± 0.43 1.47 ± 0.82 0.46 ± 0.26 DC 58 0.60 ± 0.31 1.00 ± 0.35 0.43 ± 0.17 FH 12 0.69 ± 0.28 1.37 ± 0.55 0.23 ± 0.17 ML 47 1.75 ± 0.91 2.96 ± 1.72 0.82 ± 0.65 MM-Da 139 1.29 ± 1.01 1.95 ± 1.49 0.55 ± 0.46 MP-H 35 0.83 ± 0.59 1.31 ± 0.67 0.3 ± 0.17 OP 10 1.16 ± 0.12 1.15 ± 0.54 0.31 ± 0.24 PR 17 0.81 ± 0.47 1.23 ± 0.59 0.27 ± 0.17 SG-R 11 1.49 ± 0.23 1.17 ± 0.58 0.32 ± 0.15 Other 75 1.26 ± 0.83 1.73 ± 1.03 0.64 ± 0.85 reg.(shade-intolerant) AP 66 0.81 ± 0.32 1.11 ± 0.39 0.29 ± 0.15 AQ 33 1.17 ± 0.70 1.58 ± 0.81 0.48 ± 0.3 DC 121 0.72 ± 0.24 0.96 ± 0.38 0.4 ± 0.2 FH 14 0.54 ± 0.20 1.03 ± 0.41 0.16 ± 0.1 HH 10 2.13 ± 0.77 1.99 ± 1.22 0.49 ± 0.31 ML 14 1.92 ± 1.22 3.35 ± 1.40 1.11 ± 0.8 MM-Da 164 1.42 ± 0.91 1.68 ± 1.06 0.52 ± 0.34 MP-H 72 0.83 ± 0.82 1.08 ± 0.38 0.24 ± 0.15 PR 17 0.58 ± 0.13 1.16 ± 0.38 0.31 ± 0.19 SG-R 16 0.56 ± 0.04 0.94 ± 0.29 0.36 ± 0.24 WU 12 1.76 ± 1.10 2.40 ± 1.54 0.8 ± 0.61 Others 71 1.04 ± 0.87 1.51 ± 0.85 0.4 ± 0.33 Mixed stand planted trees ML 59 21.51 ± 6.7 22.88 ± 5.1 3.64 ± 1.06 MM-Da 141 22.60 ± 8.0 21.08 ± 3.7 3.84 ± 1.31 reg.(shade-tolerant) AP 36 0.66 ± 0.29 1.28 ± 1.76 0.31 ± 0.12 DC 24 0.83 ± 0.56 1.01 ± 0.6 0.29 ± 0.18 ER 30 0.57 ± 0.27 1.03 ± 0.52 0.22 ± 0.13 LN 11 0.58 ± 0.29 1.09 ± 0.5 0.33 ± 0.15 MJ-J 12 2.11 ± 1.25 1.82 ± 1.61 0.55 ± 0.45 ML 367 1.51 ± 1.00 2.34 ± 1.32 0.66 ± 0.54 MM-Da 53 2.76 ± 3.40 2.01 ± 2.51 0.66 ± 0.79 MP-H 17 0.74 ± 0.43 0.85 ± 0.52 0.26 ± 0.17 WZⅨ 12 0.74 ± 0.40 0.97 ± 0.56 0.36 ± 0.31 Other 105 1.48 ± 1.0 1.25 ± 0.9 0.39 ± 0.43 reg.(shade-intolerant) AP 29 0.88 ± 0.54 1.31 ± 0.65 0.37 ± 0.22 DC 14 0.74 ± 0.58 0.82 ± 0.52 0.26 ± 0.1 LN 12 0.61 ± 0.42 1.17 ± 0.29 0.25 ± 0.09 ML 305 1.42 ± 0.84 2.21 ± 1.23 0.61 ± 0.46 MM-Da 47 1.85 ± 2.77 1.52 ± 1.62 0.45 ± 0.45 MP-H 20 0.48 ± 0.05 0.80 ± 0.33 0.25 ± 0.13 PR 23 0.63 ± 0.33 1.27 ± 0.45 0.4 ± 0.16 Others 79 1.61 ± 1.20 1.21 ± 0.92 0.4 ± 0.38 2.3.2 Point Pattern Analysis The pair correlation function which is based on the distance between a pair of points (PCF, \(\:g\left(r\right)=\frac{\widehat{K}\left(r\right)}{2\pi\:r}\) , \(\:\widehat{K}\left(r\right)=A\sum\:_{i=1}^{N}\sum\:_{j=1}^{N}\frac{{w}_{ij}\left(d\right)}{{N}^{2}}\) , A = area; N = number of trees; r = the observation scale (0, 1, 2… 20 m); λ = density; w ij is the weight.) can analyze tree spatial distribution patterns across scales and to identify the scales at which patterns may change, thereby facilitating the interpretation of spatial patterns (Getzin et al., 2006 ; Wiegand and Moloney, 2014 ). Therefore, we used the univariate distribution model g 11 ( r ) of PCF to analyze the spatial distributions of stands, planted trees, and regenerations. We adopted complete spatial randomness (CSR) as the null model (Wiegand and Moloney, 2014 ), and the Monte Carlo (MC) method was used to simulate the observed values 199 times. The observation scale was set to half the short side of the quadrat ( r = 0–20 m) and the envelope curves were plotted using 5% of the maximum and minimum values. Observations falling above, within, and below the simulation interval indicate significant aggregation, randomness, and regularity of the distribution pattern at the corresponding scales, respectively (Li et al., 2021b ). 2.3.3 Analysis of Spatial Correlation Plantations have a clear origin, with unified planting time and locations, high management intensity, and relatively homogeneous habitats (Bremer and Farley, 2010 ; Paquette and Messier, 2010 ), even small plots can reveal interspecific spatial correlations. Therefore, we applied the case-control method to analyze the spatial correlation between planted trees and regenerations (Wiegand and Moloney, 2004 ; Diggle et al., 2007 ), the coordinates of planted trees were fixed as the control (Pattern 1), whereas regenerations were treated as the case (Pattern 2) and were randomly distributed around the control, the results will reflect the influence of planted trees on the distribution of regenerations. We used the bivariate distribution model g 12 ( r ) to perform this analysis. In addition, using CSR as the null model, we treated the shade-tolerant group as Pattern 1 and the shade-intolerant group as Pattern 2, and analyzed spatial correlations between functional groups using g 12 ( r ) and g 21 ( r ), the parameter settings were consistent with those for the univariate distribution model g 11 ( r ). At the observation scale r , g 12 ( r ) represents the distribution of Pattern 2 around Pattern 1, i.e., the spatial correlation of Pattern 2 relative to Pattern 1. Similarly, at the observation scale r , g 21 ( r ) represents the spatial correlation of Pattern 1 relative to Pattern 2. The theoretical standard value for spatial correlation is 1, when g 12 ( r ) and g 21 ( r ) > 1, the correlation is positive, that means attraction, when g 12 ( r ) and g 21 ( r ) < 1, the correlation is negative, that means repulsion, and when g 12 ( r ) and g 21 ( r ) = 1, there is no correlation, that means independence (Getzin et al., 2006 ; Wiegand and Moloney, 2014 ). When the observed values exceeded the 95% simulation envelopes, the spatial correlation is considered significant, otherwise the relationship is regarded as weak. All analyses were conducted in Programita (version, 2018) (Wiegand and Moloney, 2014 ), and graphing were produced in R (version, 4.4.3) using the ggplot2 package. 3. Results 3.1 Spatial Distribution Patterns of M. macclurei and M. laosensis Plantations In both monoculture stands and the mixed stand showed aggregated distribution patterns, but the scales differed (Fig. 2 a- 2 c). Among them, the MM-Da Stand was aggregated at r = 0–16 m, whereas the ML Stand and the Mixed Stand were aggregated at r = 0–12 m. In the ML Stand, planted trees were aggregated at r = 0–2 m and were randomly distributed at other scales (Fig. 2 d). In the MM-Da Stand and the Mixed Stand, planted trees were aggregated at r = 0–3 m and r = 0–2 m, respectively (Fig. 2 e- 2 f). Regenerations in the ML Stand was aggregated at r = 0–14 m (Fig. 2 g), whereas regenerations in the MM-Da Stand was aggregated at almost all scales (Fig. 2 h). In the Mixed Stand, the aggregation of regenerations was similar to that in the ML Stand (Fig. 2 i). 3.2 Spatial Distribution Patterns of Functional groups In both monoculture stands, the shade-tolerant group was aggregated at all observation scales (Fig. 3 a- 3 b). In contrast, in the Mixed Stand, it was aggregated only at r = 0–8 m (Fig. 3 c). In the ML Stand, the shade-intolerant group was mainly aggregated at r = 0–13 m (Fig. 3 d). However, in the MM-Da Stand, its aggregation scale expanded to r = 0–20 m (Fig. 3 e). Moreover, the aggregation intensity of the shade-intolerant group was lower in the Mixed Stand than in monoculture stands, occurring only in the range of r = 0–7 m, while exhibiting a random distribution in the range of r = 8–16 m, and regular at the remaining scales (Fig. 3 f). 3.3 Spatial Correlations between Planted trees and Regeneration In the ML Stand and the Mixed Stand, planted trees and regeneration were positively associated at r = 0–2 m, but in the MM-Da Stand, they were positively associated at r = 0–2 m but negatively associated at other scales (Fig. 4 a- 4 c). Under both planting patterns, planted trees and the shade-tolerant group were positively associated at small scales ( r = 0–5 m), whereas they showed random correlation at other scales (Fig. 4 d- 4 f). In the ML Stand, planted trees and the shade-intolerant group were repulsive at r = 0–3 m, and they were independent at the remaining scales (Fig. 4 g). The spatial correlation between planted trees and the shade-intolerant group was negative in the MM-Da Stand (Fig. 4 h), but in the Mixed Stand, it was negative at r = 0–4 m but was mainly random at the remaining scales (Fig. 4 i). 3.4 Spatial Correlations between Functional Groups The spatial correlation between the shade-tolerant and shade-intolerant groups was negative at small scales in all stands, but their relationships at other scales differed slightly (Fig. 5 a- 5 c). Specifically, the two groups showed attraction at r = 4–20 m and r = 6–20 m in both monoculture stands, respectively. In contrast, in the Mixed Stand, they showed attraction at r = 4–10 m but repulsion at r = 12–20 m. Moreover, in both monoculture and mixed stands, the spatial correlation of the shade-intolerant group relative to the shade-tolerant group showed patterns similar to those of the shade-tolerant group relative to the shade-intolerant group (Fig. 5 d- 5 f). 4. Discussion 4.1 Plantations Exhibit Aggregated Distribution after Near-Natural Transformation Aggregated distribution is the most common spatial pattern in nature (Salas et al., 2006 ; Getzin et al., 2008 ; Wiegand et al., 2017 ), and is also an important trend during plantation succession (Li et al., 2021c ; Yang et al., 2025 ). Other studies have shown that plantation spatial patterns are influenced not only by ecological processes such as regeneration, self-thinning, and external disturbances, but also jointly regulated by abiotic conditions including topography, soil, and light conditions (Li et al., 2021a ; Matsushita et al., 2025 ), thereby shifting toward non-regular patterns, which is consistent with our results. In the M. macclurei and M. laosensis plantations, planted trees show aggregation at small scales (Appendix 1), and an important reason is the occurrence of sprouting (Yang et al., 2025 ), which supports Hypothesis 1. Sprouting is a strategy by many woody plants to maintain their niches under competition or disturbance, and its occurrence is closely related to species functional traits (Bond and Midgley, 2001 ; Clarke et al., 2013 ). The drivers of sprouting may differ among ecosystems and species (Pausas et al., 2016 ), and our understanding of how sprouting occurs in planted trees and what factors influence it in plantations remains limited (Shi et al., 2025 ). However, it is certain that plantation species that undergo strong differentiation at the root during the first planting are rare. Competition drives the mortality of some trees (Zhang et al., 2020 ), but the extent of pattern change is also influenced by differences in functional traits of planted trees (Forrester et al., 2018 ; Zambrano et al., 2020 ), and these complex relationships further cause the spatial patterns of planted trees to deviate from regularity. In Pingxiang, which is also in the southern subtropical region, the spatial pattern of planted trees in a Chinese fir mixed plantation shifts from a regular pattern at establishment to an aggregated pattern after 27 years of succession (Li et al. 2021a , c ), supporting our results. Changes in plantation spatial patterns are closely related to planting design, species composition, and management duration. The aggregated distribution of regeneration precisely supports Hypothesis 1. Both planted trees have reached physiological maturity and continuously provide seed sources for regeneration, and their capsule and aggregate-fruit traits further cause most seeds to fall near parent trees (Appendix 1). Accordingly, regeneration occurs mainly within the neighborhood of planted trees and shows an aggregated pattern (Appendix 2). Dispersal limitation is also considered an important mechanism underlying aggregated patterns in natural forests (Condit et al., 2000 ; Seidler and Plotkin, 2006 ; Szwagrzyk et al., 2021 ). Because plantations typically have clear management rotations and planted trees are often clear-cut at maturity, regeneration following seed production has been less studied (Yang et al., 2025 ). The soil seed bank is another important source of regeneration (Nathan and Muller-Landau, 2000 ; Singh et al., 2023 ), and it may also increase the intensity and spatial scale of regeneration aggregation. In addition, seedlings are highly sensitive and dependent on microhabitat conditions such as light, moisture, soil, and topography (De Lombaerde et al., 2020 ; Sangsupan et al., 2021 ). Environmental filtering therefore promotes regeneration establishment in locally favorable sites, resulting in aggregation. Both shade-tolerant and shade-intolerant regeneration groups show aggregated distributions, but their aggregation intensity differs between monoculture and mixed stands. This pattern is widely considered to be related to understory light conditions (Valladares and Niinemets, 2008 ; Klopčič et al., 2015 ). Row planting creates relatively regular canopy gaps in the stand, with alternating distribution of high and low light in the understory, resulting in a patchy light environment (Zhang et al., 2019 ; Schneider et al., 2021 ). Such uneven light conditions cause shade-tolerant and shade-intolerant regenerations to occur in different light environments. Because M. laosensis has broad leaves and a dense canopy, shade-intolerant regeneration tends to occur in canopy gaps in its monoculture stand. In contrast, M. macclurei has smaller leaves and a relatively large crown, and its monoculture stand has better light conditions. As a result, both functional groups can establish under the canopy or in gaps (Yang et al., 2025 ). In the Mixed Stand, differences in crown structure between planted trees create vertical stratification and increase within-stand light availability (Pretzsch, 2014 ; Forrester et al., 2018 ). Consequently, both shade-tolerant and shade-intolerant groups can establish in multiple positions, which reduces their aggregation scale and intensity (Yang et al., 2025 ). Overall, planted trees, planting patterns, and spatial configuration influence the occurrence and distribution patterns of regeneration by altering canopy structure and microhabitat conditions. 4.2 Spatial Correlations Between Planted Trees and Regenerations in Plantations The degree to which regenerations depend on light determines the direction and strength of their association with planted trees, supporting Hypothesis 2. Shade-tolerant groups can maintain growth under low-light conditions, and their regeneration processes are less dependent on forest gaps (Montgomery and Chazdon, 2001 ; Klopčič et al., 2015 ). Therefore, they can utilize the relatively stable light environment beneath the canopies of planted trees. In contrast, shade-intolerant regenerations tend to occur away from planted trees. For example, in Pinus massoniana plantations, high planting density and canopy closure place seedlings under low-light conditions, which reduces their photosynthetic potential and affects growth (Yan et al., 2019 ; Jin et al., 2024 ; Hao et al., 2025 ). Accordingly, during the near-natural transformation of plantations, the light requirements of seedlings can be leveraged by supplementing shade-tolerant seedlings under the canopy or thinning to improve within-stand light conditions (Cheng et al., 2017 ; Su et al., 2024 ), thereby promoting regeneration and growth within forest gaps and accelerating the transition of plantations from single-layer to multi-layered mixed structures (Liu et al., 2018 ). However, some tree species adopt flexible light adaptation strategies during growth. For instance, Michelia macclurei exhibits strong shade-tolerance during the seedling stage, enabling survival in forest gaps and under tree canopies, while becoming light-preferring in later developmental stages (Li et al., 2021c ; Yang et al., 2025 ). Therefore, when both shade-tolerant and shade-intolerant groups are considered at the stand level, the spatial correlation between planted trees and regenerations may be diluted (Fig. 4 a- 4 c). This also explains why some studies have found that spatial correlations between planted trees and regenerations are not characterized by a single pattern, with its manifestation varying according to stand structure, observation scale, and functional group differentiation (Ali et al., 2019 ; Li et al., 2021c ; Zhang et al., 2022 ). 4.3 Scale-dependent Spatial Correlations between Functional Groups in Plantations Light resource partitioning and habitat preferences segregate shade-tolerant and shade-intolerant groups (Montgomery and Chazdon, 2001 ; Valladares and Niinemets, 2008 ; Bartkowicz and Paluch, 2019 ), exhibiting negative correlations at small scales (Fig. 5 a- 5 c). However, as the observation scale increases, their spatial correlations are dominated by positive correlations, reflecting the mosaic distribution of light patches in plantations. After the distributions of functional groups span both canopy and gap scales, they tend toward complementary coexistence. This distribution pattern differs from interspecific correlations in natural forests, which often become random as the observation scale increases (Getzin et al., 2006 ; Wiegand et al., 2007 ; Cavard et al., 2011 ). Consistently, in a forest-edge planting experiment in southern Sweden (Wiström and Nielsen, 2014 ), they found that vegetation composed of planted trees and regeneration with different shade tolerance showed spatial differentiation along the forest edge 16 years after establishment. Light-demanding pioneer species occupied high-light environments, whereas shade-tolerant trees and shrubs primarily distributed under relatively low-light canopies, strongly supporting our findings. However, spatial correlations between functional groups may be transient, and planting patterns influence future shifts in functional group spatial correlations (Li et al., 2021c ). In monoculture stands, microhabitat types are limited and functional-group differentiation is constrained (Pretzsch and Zenner, 2017 ), potentially maintaining spatial associations over extended periods. In mixed stands, interactions among multiple species are alleviated (Forrester, 2014 ; Li et al., 2021c ), may weaken the associations. Furthermore, spatial correlations between functional groups may be regulated by belowground components. Plant-soil feedbacks (PSFs) promote plant coexistence by regulating relative competitive advantages of species in high-light and low-light environments (Xi et al., 2023 ), thereby reducing correlation strength. This suggests that spatial correlations between functional groups in plantations exhibit spatiotemporal effects and are influenced by multiple ecological processes. 5. Conclusion Following the transition of plantations toward near-natural transformation, their spatial patterns have undergone significant alterations. The point patterns of trees in M. laosensis and M. macclurei plantations deviate from the initial planting patterns. The distribution patterns of these plantations exhibit aggregated distributions, driven jointly by sprouting, seed dispersal limitation, and within-stand light conditions. This suggests that spatial patterns are controlled by multiple ecological processes, with aggregation represents the ultimate spatial form of forest communities. Tree functional traits determine the direction of spatial correlations between planted trees and regenerations. Therefore, the selection of planted species and planting patterns may be key to maintaining stable regeneration in plantations. Because our study includes only two major local afforestation species, the application of additional species and functional trait combinations in near-natural transformation requires further research. Shade-tolerant groups mainly occur beneath canopies, whereas shade-intolerant groups prefer canopy gaps. Light conditions largely determine patterns of interspecific association, making seedling shade tolerance a priority consideration in the near-natural transformation of plantations. Improving interspecific relationships contributes to optimizing stand structure and enhancing functional stability. The spatial patterns and spatial correlations observed in M. laosensis and M. macclurei plantations deepen our understanding of near-natural transformation processes, and they may also be applicable to forest management practice. Declarations Ethics approval and consent to participate: Not applicable Consent for publication: Not applicable Competing interests: Authors have no conflicts of interest Funding: This work was supported the National Natural Science Foundation of China (Grant No. 32060340). Author Contribution Yuanfa Li conceived the idea, Ziyi Chen and Yuanfa Li wrote the manuscript, all authors collected dataset from wild and reviewed the manuscript. Acknowledgement Master Studies from College of Forestry, Guangxi University investigated dataset in wild, they were Jintao Li, Yongyi Wei, Liting Wei, Xiuqiong Luo. Data Availability The datasets used and/or analysed during the current study are available from the corresponding author on reasonable request. References Ali A, Dai D, Akhtar K, Teng M, Yan Z, Urbina-Cardona N, Mullerova J, Zhou Z (2019) Response of understory vegetation, tree regeneration, and soil quality to manipulated stand density in a Pinus massoniana plantation. Global Ecology and Conservation 20: e00775.https://doi.org/10.1016/j.gecco.2019.e00775 Asanok L, Marod D, Duengkae P, Pranmongkol U, Kurokawa H, Aiba M, Katabuchi M, Nakashizuka T (2013) Relationships between functional traits and the ability of forest tree species to reestablish in secondary forest and enrichment plantations in the uplands of northern Thailand. 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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-8701430","acceptedTermsAndConditions":true,"allowDirectSubmit":false,"archivedVersions":[],"articleType":"Research Article","associatedPublications":[],"authors":[{"id":618388698,"identity":"066ffbd7-9de4-43bd-a319-4c769652d63e","order_by":0,"name":"Ziyi Chen","email":"","orcid":"","institution":"Guangxi University","correspondingAuthor":false,"prefix":"","firstName":"Ziyi","middleName":"","lastName":"Chen","suffix":""},{"id":618388701,"identity":"006c3ac2-b3e8-4b7a-b735-5a94cee7b6a4","order_by":1,"name":"Liuqi Feng","email":"","orcid":"","institution":"Guangxi University","correspondingAuthor":false,"prefix":"","firstName":"Liuqi","middleName":"","lastName":"Feng","suffix":""},{"id":618388702,"identity":"f90da8ec-c8ed-4eaa-8e12-7000f55f1022","order_by":2,"name":"Yifan Liu","email":"","orcid":"","institution":"Guangxi University","correspondingAuthor":false,"prefix":"","firstName":"Yifan","middleName":"","lastName":"Liu","suffix":""},{"id":618388705,"identity":"6d5e63a1-f398-4583-b664-09994aecc8ef","order_by":3,"name":"Yuanfa Li","email":"data:image/png;base64,iVBORw0KGgoAAAANSUhEUgAAAZAAAAAyAQMAAABI0h/eAAAABlBMVEX///8AAABVwtN+AAAACXBIWXMAAA7EAAAOxAGVKw4bAAAAxUlEQVRIiWNgGAWjYBACxv7mgw8kKv7J8ROthXnGsWQDizMHjCUbiNXC3pBjJlHZciBxwwFitfA2HDCTuNlwJ3Hz8eQNDD8qthHWItnckGw5c8cz421nnhUw9py5TViLYcOBg7clzzDLbruRY8DM2EaEFvsDiQ3Sf9uYGTfPIFYLY0Myk4Rk22HFDRJEa5lxjNlA4kyasQTQLweJ8gtjf/9HYFTayPG3J2988KOCCC1IIMHgAEnqwVpI1TEKRsEoGAUjBAAAz7NGWrDf8XoAAAAASUVORK5CYII=","orcid":"","institution":"Guangxi University","correspondingAuthor":true,"prefix":"","firstName":"Yuanfa","middleName":"","lastName":"Li","suffix":""}],"badges":[],"createdAt":"2026-01-26 14:39:31","currentVersionCode":1,"declarations":"","doi":"10.21203/rs.3.rs-8701430/v1","doiUrl":"https://doi.org/10.21203/rs.3.rs-8701430/v1","draftVersion":[],"editorialEvents":[],"editorialNote":"","failedWorkflow":false,"files":[{"id":106727315,"identity":"3389cc8f-122a-4ca9-a0c0-12cc6739c4ba","added_by":"auto","created_at":"2026-04-12 18:38:40","extension":"jpg","order_by":1,"title":"Figure 1","display":"","copyAsset":false,"role":"figure","size":4322379,"visible":true,"origin":"","legend":"\u003cp\u003eThe map shows the locations of pure and mixed forests. The dots represent tree species distribution, with red representing planted trees, blue representing shade-tolerant regenerations, and yellow representing shade-intolerant regenerations.\u003c/p\u003e","description":"","filename":"Fig.1location.jpg","url":"https://assets-eu.researchsquare.com/files/rs-8701430/v1/42184d462241684fe7f1c1ca.jpg"},{"id":106959626,"identity":"ef66b7af-ceca-4629-91b7-88d9e18003d2","added_by":"auto","created_at":"2026-04-15 09:12:26","extension":"jpg","order_by":2,"title":"Figure 2","display":"","copyAsset":false,"role":"figure","size":795363,"visible":true,"origin":"","legend":"\u003cp\u003ePoint distribution patterns of ML Stand, MM-Da Stand, and \u003cem\u003eM. laosensis\u003c/em\u003e -\u003cem\u003e M. macclurei\u003c/em\u003e Mixed Stand, as well as the point distribution patterns of planted trees and regeneration based on the completely spatial random (CSR) null model. The gray area represents the 95% Monte Carlo simulation area, the black solid line marked with red dots is the observed value, and the red dashed line is the expected value 1. n represents the number of individuals.\u003c/p\u003e","description":"","filename":"Fig.2pointdistributionpatternsofstands.jpg","url":"https://assets-eu.researchsquare.com/files/rs-8701430/v1/da58524277854195f422cc9a.jpg"},{"id":106645068,"identity":"97ca6e12-23a5-4461-a4b7-1e72b1e096b6","added_by":"auto","created_at":"2026-04-10 19:52:55","extension":"jpg","order_by":3,"title":"Figure 3","display":"","copyAsset":false,"role":"figure","size":565136,"visible":true,"origin":"","legend":"\u003cp\u003ePoint distribution patterns of shade-tolerant and shade-intolerant functional groups in ML Stand, MM-Da Stand, and \u003cem\u003eM. laosensis\u003c/em\u003e -\u003cem\u003e M. macclurei\u003c/em\u003e Mixed Stand. The gray area represents the 95% Monte Carlo simulation area, the black solid line marked with red dots is the observed value, and the red dashed line is the expected value 1. n represents the number of individuals.\u003c/p\u003e","description":"","filename":"Fig.3pointdistributionpatternsoffunctionalgroups.jpg","url":"https://assets-eu.researchsquare.com/files/rs-8701430/v1/2160cfa8e2c378710e164aa4.jpg"},{"id":107479257,"identity":"a5b148e1-eae5-4b14-9d9f-a53467236835","added_by":"auto","created_at":"2026-04-22 01:21:22","extension":"jpg","order_by":4,"title":"Figure 4","display":"","copyAsset":false,"role":"figure","size":839146,"visible":true,"origin":"","legend":"\u003cp\u003eSpatial correlation of planted trees and regenerations in ML Stand, MM-Da Stand, and \u003cem\u003eM. laosensis\u003c/em\u003e -\u003cem\u003e M. macclurei\u003c/em\u003e Mixed Stand. The gray area represents the 95% Monte Carlo simulation area, the black solid line marked with red dots is the observed value, and the red dashed line is the expected value 1.\u003c/p\u003e","description":"","filename":"Fig.4spatialcorrelationofplantedandregeneration.jpg","url":"https://assets-eu.researchsquare.com/files/rs-8701430/v1/b4a4d103673c778178214539.jpg"},{"id":106645070,"identity":"04134869-dc56-4ae2-a60b-81b32dd35306","added_by":"auto","created_at":"2026-04-10 19:52:55","extension":"jpg","order_by":5,"title":"Figure 5","display":"","copyAsset":false,"role":"figure","size":570527,"visible":true,"origin":"","legend":"\u003cp\u003eSpatial correlation between shade-tolerant and shade-intolerant functional groups in ML Stand, MM-Da Stand, and \u003cem\u003eM. laosensis\u003c/em\u003e -\u003cem\u003e M. macclurei\u003c/em\u003e Mixed Stand based on CSR. The gray area represents the 95% Monte Carlo simulation area, the black solid line marked with red dots is the observed value, and the red dashed line is the expected value 1.\u003c/p\u003e","description":"","filename":"Fig.5spatialcorrelationbetweenfunctionalgroups.jpg","url":"https://assets-eu.researchsquare.com/files/rs-8701430/v1/013a5bdc7e9f22a7a8ac9e68.jpg"},{"id":107479262,"identity":"ac84deec-a6d0-423c-b9cf-0bddfce6ddee","added_by":"auto","created_at":"2026-04-22 01:21:30","extension":"pdf","order_by":0,"title":"","display":"","copyAsset":false,"role":"manuscript-pdf","size":7807242,"visible":true,"origin":"","legend":"","description":"","filename":"manuscript.pdf","url":"https://assets-eu.researchsquare.com/files/rs-8701430/v1/efea3e75-327b-4452-aa21-b2fa5f4680e9.pdf"}],"financialInterests":"No competing interests reported.","formattedTitle":"Spatial Patterns of Michelia macclurei and Mytilaria laosensis plantations after 24 Years of Near-Natural Transformation","fulltext":[{"header":"1. Introduction","content":"\u003cp\u003eSpatial patterns serve as a crucial method for investigating the relationship between community structure and ecological processes, with their core encompassing the spatial distribution of individuals and interspecies correlations (Getzin et al., \u003cspan citationid=\"CR37\" class=\"CitationRef\"\u003e2008\u003c/span\u003e). Distribution patterns reveal the degree of aggregation within communities across multiple scales through the distribution of tree points, reflecting the effects of seed dispersal limitations, heterogeneous resource availability, and historical disturbances and succession (Pommerening, \u003cspan citationid=\"CR89\" class=\"CitationRef\"\u003e2002\u003c/span\u003e). Spatial correlations describe the spatial relationships among individuals from different populations, life stages, or origins, providing important evidence for interpreting the maintenance of species diversity, niche differentiation, and community stability (Getzin et al., \u003cspan citationid=\"CR37\" class=\"CitationRef\"\u003e2008\u003c/span\u003e; Ben-Said, \u003cspan citationid=\"CR7\" class=\"CitationRef\"\u003e2021\u003c/span\u003e). Spatial patterns have been extensively used to elucidate the mechanisms of species diversity formation and coexistence in natural forests (Stoll and Newbery, \u003cspan citationid=\"CR113\" class=\"CitationRef\"\u003e2005\u003c/span\u003e; Gu et al., \u003cspan citationid=\"CR39\" class=\"CitationRef\"\u003e2019\u003c/span\u003e), but rarely used to analyze the succession and regeneration of spatial structure in plantations (Ben-Said, \u003cspan citationid=\"CR7\" class=\"CitationRef\"\u003e2021\u003c/span\u003e). However, with the advancement of ecological civilization construction and the deepening of the concept of multifunctional forest utilization, the management of monoculture forests guided by maximizing timber production and economic benefits have gradually exposed problems such as insufficient ecosystem service functions (Paquette and Messier, \u003cspan citationid=\"CR85\" class=\"CitationRef\"\u003e2010\u003c/span\u003e; Liu et al., \u003cspan citationid=\"CR63\" class=\"CitationRef\"\u003e2018\u003c/span\u003e). Many stands are undergoing near-natural transformation in order to adjust stand structure and enhance ecosystem functions (Ming et al., \u003cspan citationid=\"CR73\" class=\"CitationRef\"\u003e2020\u003c/span\u003e; O\u0026rsquo;Hara, \u003cspan citationid=\"CR81\" class=\"CitationRef\"\u003e2016\u003c/span\u003e), yet the transformation process and underlying drivers remain unclear. Applying spatial pattern analysis to plantations research helps deepen our understanding of community structural dynamics and provides scientific support for optimizing management strategies (Forrester, \u003cspan citationid=\"CR31\" class=\"CitationRef\"\u003e2014\u003c/span\u003e; Zhang et al., \u003cspan citationid=\"CR140\" class=\"CitationRef\"\u003e2019\u003c/span\u003e).\u003c/p\u003e \u003cp\u003ePlanting patterns, namely monoculture stands and mixed stand, influence the spatial patterns of plantations under natural succession. The species composition, spatial arrangement, and proportions of planted trees directly determine the intensity of neighborhood interactions (Forrester, \u003cspan citationid=\"CR31\" class=\"CitationRef\"\u003e2014\u003c/span\u003e; Pretzsch and Sch\u0026uuml;tze, \u003cspan citationid=\"CR93\" class=\"CitationRef\"\u003e2021\u003c/span\u003e). Individual plants in monoculture forests have similar resource requirements, and the intense competition within their neighborhoods makes them prone to self-thinning (Hui et al., \u003cspan citationid=\"CR47\" class=\"CitationRef\"\u003e2018\u003c/span\u003e), which alters the spatial patterns. In contrast, differences in growth processes and resource utilization among species in mixed stand may alleviate or alter interaction ways between neighboring species (Forrester, \u003cspan citationid=\"CR31\" class=\"CitationRef\"\u003e2014\u003c/span\u003e; Pretzsch and Sch\u0026uuml;tze, \u003cspan citationid=\"CR93\" class=\"CitationRef\"\u003e2021\u003c/span\u003e). Meanwhile, planting patterns affect seed composition and sources (Cole et al., \u003cspan citationid=\"CR19\" class=\"CitationRef\"\u003e2010\u003c/span\u003e; Liu et al., \u003cspan citationid=\"CR65\" class=\"CitationRef\"\u003e2022\u003c/span\u003e), thereby altering the establishment processes and spatial distribution patterns of regenerations (Huanca Nu\u0026ntilde;ez et al., \u003cspan citationid=\"CR43\" class=\"CitationRef\"\u003e2021\u003c/span\u003e; Wang et al., \u003cspan citationid=\"CR121\" class=\"CitationRef\"\u003e2019\u003c/span\u003e). In monoculture stands, parent trees are dominated by a single species, and regenerations mainly originates from seed rain of planted trees and soil seed banks (Seidler and Plotkin, \u003cspan citationid=\"CR107\" class=\"CitationRef\"\u003e2006\u003c/span\u003e; Liu et al., \u003cspan citationid=\"CR65\" class=\"CitationRef\"\u003e2022\u003c/span\u003e). By contrast, parent trees in mixed stand are more diverse, expanding seed sources and dispersal range, better regulating the forest microclimate, increasing the quantity and quality of litter (Huang et al., \u003cspan citationid=\"CR45\" class=\"CitationRef\"\u003e2017\u003c/span\u003e; Zhang et al., \u003cspan citationid=\"CR142\" class=\"CitationRef\"\u003e2022\u003c/span\u003e), and promoting the regeneration of site-quality-sensitive species (De Frenne et al., \u003cspan citationid=\"CR23\" class=\"CitationRef\"\u003e2019\u003c/span\u003e). Furthermore, due to complex structure, mixed stand shows self-regulation and resistance to disturbance during succession (Liu et al., \u003cspan citationid=\"CR63\" class=\"CitationRef\"\u003e2018\u003c/span\u003e; Pretzsch and Zenner, \u003cspan citationid=\"CR95\" class=\"CitationRef\"\u003e2017\u003c/span\u003e), which increases the differences in spatial structure between mixed and monoculture stands.\u003c/p\u003e \u003cp\u003eIn addition to planting patterns, the functional traits of regeneration also influence the spatial patterns of plantations (Asanok et al., \u003cspan citationid=\"CR3\" class=\"CitationRef\"\u003e2013\u003c/span\u003e; Rahayu et al., \u003cspan citationid=\"CR97\" class=\"CitationRef\"\u003e2022\u003c/span\u003e). Specifically, differences among functional groups in resource acquisition and utilization affect regeneration processes and further modify spatial patterns (Forrester, \u003cspan citationid=\"CR31\" class=\"CitationRef\"\u003e2014\u003c/span\u003e; Zambrano et al., \u003cspan citationid=\"CR138\" class=\"CitationRef\"\u003e2020\u003c/span\u003e). Although many factors influence the renewal process, light may be the most important limiting factor at small spatial scales (De Lombaerde et al., \u003cspan citationid=\"CR25\" class=\"CitationRef\"\u003e2020\u003c/span\u003e; Sangsupan et al., \u003cspan citationid=\"CR103\" class=\"CitationRef\"\u003e2021\u003c/span\u003e). Shade-tolerant groups have lower light compensation points and higher light-use efficiency (Valladares and Niinemets, \u003cspan citationid=\"CR119\" class=\"CitationRef\"\u003e2008\u003c/span\u003e), which enables them to maintain photosynthesis and growth under low understory light. Therefore, they readily form small-scale aggregations beneath or near the canopy of planted trees (Getzin et al., \u003cspan citationid=\"CR37\" class=\"CitationRef\"\u003e2008\u003c/span\u003e; Paluch, \u003cspan citationid=\"CR83\" class=\"CitationRef\"\u003e2024\u003c/span\u003e). In contrast, shade-intolerant groups are highly dependent on light, prefer canopy gaps or forest edges, and their spatial patterns are closer to dispersed or random distributions (Klopčič et al., \u003cspan citationid=\"CR51\" class=\"CitationRef\"\u003e2015\u003c/span\u003e). Current research on plantations mainly examines regeneration dynamics from the perspectives of species composition and environmental drivers, focusing on the effects of habitat heterogeneity and competitive pressure (Ali et al., \u003cspan citationid=\"CR1\" class=\"CitationRef\"\u003e2019\u003c/span\u003e; Kremer et al., \u003cspan citationid=\"CR53\" class=\"CitationRef\"\u003e2022\u003c/span\u003e). However, few studies analyze the role of regeneration functional groups in shifts in stand spatial patterns.\u003c/p\u003e \u003cp\u003eThe subtropical region of southern China boasts favorable hydrothermal conditions and numerous native tree species occur, which play important roles in regional ecological restoration, timber production, and landscape construction (Lu et al., \u003cspan citationid=\"CR69\" class=\"CitationRef\"\u003e2017\u003c/span\u003e; Li et al., \u003cspan citationid=\"CR59\" class=\"CitationRef\"\u003e2021c\u003c/span\u003e). In some areas, monoculture and mixed stands of native species have been established for decades, but most stands were harvested once the planted trees reach economic maturity. Using \u003cem\u003eMytilaria laosensis\u003c/em\u003e Lecomte and \u003cem\u003eMichelia macclurei\u003c/em\u003e Dandy stands that have exceeded one rotation period (20 a) as examples, this study examines how planting patterns influence the spatial patterns of native-species plantations and hypothesizes: (1) Mortality of planted trees and the occurrence of regeneration (Zhang et al., \u003cspan citationid=\"CR140\" class=\"CitationRef\"\u003e2019\u003c/span\u003e; Li et al., \u003cspan citationid=\"CR59\" class=\"CitationRef\"\u003e2021c\u003c/span\u003e) shift the stand point pattern from an initially regular distribution to a random or aggregated distribution. Moreover, seed dispersal modes and dispersal limitation affect the distribution of regeneration (Nathan and Muller-Landau, \u003cspan citationid=\"CR77\" class=\"CitationRef\"\u003e2000\u003c/span\u003e; Dent and Estrada-Villegas, \u003cspan citationid=\"CR27\" class=\"CitationRef\"\u003e2021\u003c/span\u003e), leading to aggregation (Hypothesis 1); (2) Regeneration establishment depends strongly on the overstory vegetation and the microenvironment it creates (O\u0026rsquo;Brien et al., \u003cspan citationid=\"CR79\" class=\"CitationRef\"\u003e2007\u003c/span\u003e; Rautio et al., \u003cspan citationid=\"CR99\" class=\"CitationRef\"\u003e2023\u003c/span\u003e), therefore planted trees and regenerations attract each other spatially (Hypothesis 2); (3) Differences in the plant morphology and vertical structure of planted trees between monoculture and mixed stands (Montgomery and Chazdon, \u003cspan citationid=\"CR75\" class=\"CitationRef\"\u003e2001\u003c/span\u003e; Pretzsch, \u003cspan citationid=\"CR91\" class=\"CitationRef\"\u003e2014\u003c/span\u003e), cause functional groups with different light dependence to occupy different spatial positions (Ligot et al., \u003cspan citationid=\"CR61\" class=\"CitationRef\"\u003e2014\u003c/span\u003e; Forrester et al., \u003cspan citationid=\"CR33\" class=\"CitationRef\"\u003e2018\u003c/span\u003e), resulting in negative correlations between functional groups (Hypothesis 3). To test these hypotheses, we established permanent plots in a \u003cem\u003eMytilaria laosensis\u003c/em\u003e monoculture stand (ML stand), a \u003cem\u003eMichelia macclurei\u003c/em\u003e monoculture stand (MM-Da stand), and \u003cem\u003eM. laosensis\u003c/em\u003e \u0026ndash; \u003cem\u003eM. macclurei\u003c/em\u003e mixed stand (Mixed stand) in southern Guangxi, and used census data for validation. The results will elucidate the drivers of change in spatial patterns and the regeneration mechanisms of functional groups, thereby promoting near-natural transformation of plantations.\u003c/p\u003e"},{"header":"2. Materials and Methods","content":"\u003cdiv id=\"Sec3\" class=\"Section2\"\u003e \u003ch2\u003e2.1 Regional Overview\u003c/h2\u003e \u003cp\u003eThe experimental site is located at the Jiepai substation of Gaofeng Forest Farm, Guangxi, southern China (108\u0026deg;21\u0026prime;\u0026ndash;108\u0026deg;24\u0026prime; E, 22\u0026deg;56\u0026prime;\u0026ndash;24\u0026deg;00\u0026prime; N). The region has a warm climate with abundant rainfall and sufficient sunshine, and it is characterized as a southern subtropical monsoon climate. The average annual temperature is 21.56\u0026deg;C, the annual precipitation is approximately 2056 mm, and the average annual sunshine duration is 1551 h. The landform is dominated by low mountains and hills with gentle relief. The parent rock is mainly granite, and long-term weathering has resulted in lateritic red soils and purple soils, with an average soil depth exceeding 80 cm and good water and fertilizer retention properties. Native tree species in the study area primarily form the overstory, including \u003cem\u003eMytilaria laosensis\u003c/em\u003e Lecomte, \u003cem\u003eMachilus macclurei\u003c/em\u003e Dandy, and \u003cem\u003eCastanopsis hystrix\u003c/em\u003e Hook. f. and Thomson ex A. DC., among others. In addition, the understory species composition is diverse, common shrubs include \u003cem\u003eMelicope pteleifolia\u003c/em\u003e (Champ. ex Benth.) Hartley, \u003cem\u003eArdisia quinquegona\u003c/em\u003e Blume, \u003cem\u003eWendlandia uvariifolia\u003c/em\u003e Hance, \u003cem\u003eActinodaphne pilosa\u003c/em\u003e (Lour.) Merr., \u003cem\u003eDesmos chinensis\u003c/em\u003e Lour., \u003cem\u003eLonicera confusa\u003c/em\u003e (Sweet) DC., and \u003cem\u003ePsychotria asiatica\u003c/em\u003e L., while common herbaceous species include \u003cem\u003eSarcandra glabra\u003c/em\u003e (Thunb.) Nakai, \u003cem\u003eSarcophyton taiwanianum\u003c/em\u003e (Hayata) Garay, \u003cem\u003eMiscanthus floridulus\u003c/em\u003e (Lab.) Warb. ex Schum. et Laut., and \u003cem\u003eDicranopteris dichotoma\u003c/em\u003e (Thunb.) Bernh, among others (Yang et al., \u003cspan citationid=\"CR136\" class=\"CitationRef\"\u003e2025\u003c/span\u003e).\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec4\" class=\"Section2\"\u003e \u003ch2\u003e2.2 Plots establishment and Data collection\u003c/h2\u003e \u003cp\u003eThe specific experimental site was located on a gentle slope with an inclination of approximately 25\u0026deg;. In 1999, we clear-cut a mature Chinese fir (\u003cem\u003eCunninghamia lanceolata\u003c/em\u003e (Lamb.) Hook.) plantation and then established monoculture stands of \u003cem\u003eMytilaria laosensis\u003c/em\u003e, \u003cem\u003eMichelia macclurei\u003c/em\u003e, and their mixed stand, with an approximate spacing of 3.5 m \u0026times; 3.5 m. The stem ratio of \u003cem\u003eMytilaria laosensis\u003c/em\u003e to \u003cem\u003eMichelia macclurei\u003c/em\u003e in the mixed stand was approximately 2:1. Manual tending was conducted during the first 1\u0026ndash;5 years after planting, after which the stands were in a state of natural succession and gradually developed a multispecies coexistence pattern (Yang et al., \u003cspan citationid=\"CR136\" class=\"CitationRef\"\u003e2025\u003c/span\u003e). In 2024\u0026ndash;2025, we established three permanent plots with sizes of 140 m \u0026times; 40 m, 70 m \u0026times; 50 m, and 70 m \u0026times; 40 m, according to the shape and area of each class (Fig.\u0026nbsp;\u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e1\u003c/span\u003e). First, we used a total station (model NTS-372R\u003csub\u003e10\u003c/sub\u003e, South Surveying and Mapping Company, Guangzhou, China) to define plot boundaries and divided each plot into n subplots of 10 m \u0026times; 10 m. Then, we mapped the coordinates (\u003cem\u003ex\u003c/em\u003e, \u003cem\u003ey\u003c/em\u003e, \u003cem\u003ez\u003c/em\u003e) of all trees within each plot (diameter at breast height, DBH\u0026thinsp;\u0026ge;\u0026thinsp;1.0 cm; DBH\u0026thinsp;\u0026lt;\u0026thinsp;1.0 cm and height\u0026thinsp;\u0026ge;\u0026thinsp;0.5 m). Finally, each tree was tagged to identify the species and to measure its dimensions. The measured variables included DBH (m), tree height (m), crown width (m), and ground diameter (cm) for non-planted species with DBH\u0026thinsp;\u0026ge;\u0026thinsp;1.0 cm or DBH\u0026thinsp;\u0026lt;\u0026thinsp;1.0 cm and height\u0026thinsp;\u0026ge;\u0026thinsp;0.5 m, these individuals included almost all planted trees and regenerations in the plots (Yang et al., \u003cspan citationid=\"CR136\" class=\"CitationRef\"\u003e2025\u003c/span\u003e).\u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec5\" class=\"Section2\"\u003e \u003ch2\u003e2.3 Data Analysis\u003c/h2\u003e \u003cdiv id=\"Sec6\" class=\"Section3\"\u003e \u003ch2\u003e2.3.1 Classification of Tree Groups\u003c/h2\u003e \u003cp\u003eTo better analyze spatial patterns in monoculture and mixed plantations and the regeneration characteristics of functional groups, we classified trees into planted trees and regenerations based on their origin, and further divided regenerations into shade-tolerant and shade-intolerant functional groups based on their light adaptability. When counting plants, sprouted individuals of planted trees below a height of 1.3 m were counted as independent plants. In ML Stand, we recorded a total of 889 individuals, including 488 planted trees, 219 shade-tolerant regenerations, and 182 shade-intolerant regenerations. In MM-Da Stand, we recorded 1318 individuals in total, including 208 planted trees, 500 shade-tolerant regenerations, and 610 shade-intolerant regenerations. In Mixed Stand, we recorded 1397 individuals, including 201 planted trees, 667 shade-tolerant regenerations, and 529 shade-intolerant regenerations (Table\u0026nbsp;\u003cspan refid=\"Tab1\" class=\"InternalRef\"\u003e1\u003c/span\u003e).\u003c/p\u003e \u003cp\u003e \u003cdiv class=\"gridtable\"\u003e\u003ctable float=\"Yes\" id=\"Tab1\" border=\"1\"\u003e \u003ccaption language=\"En\"\u003e \u003cdiv class=\"CaptionNumber\"\u003eTable 1\u003c/div\u003e \u003cdiv class=\"CaptionContent\"\u003e \u003cp\u003eBasic information on tree populations in ML stand, MM-Da stand, ML and MM-Da mixed stand. \u003cem\u003eOthers\u003c/em\u003e refer to populations with N\u0026thinsp;\u0026lt;\u0026thinsp;10. N, number of individuals; DBH, diameter at breast height; TH, tree height; CW, crown width.\u003c/p\u003e \u003c/div\u003e \u003c/caption\u003e \u003ccolgroup cols=\"7\"\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c1\" colnum=\"1\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c2\" colnum=\"2\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c3\" colnum=\"3\"\u003e\u003c/div\u003e \u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c4\" colnum=\"4\"\u003e\u003c/div\u003e \u003cdiv align=\"char\" char=\"\u0026plusmn;\" class=\"colspec\" colname=\"c5\" colnum=\"5\"\u003e\u003c/div\u003e \u003cdiv align=\"char\" char=\"\u0026plusmn;\" class=\"colspec\" colname=\"c6\" colnum=\"6\"\u003e\u003c/div\u003e \u003cdiv align=\"char\" char=\"\u0026plusmn;\" class=\"colspec\" colname=\"c7\" colnum=\"7\"\u003e\u003c/div\u003e \u003cthead\u003e \u003ctr\u003e \u003cth align=\"left\" colname=\"c1\"\u003e \u003cp\u003ePlot\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c2\"\u003e \u003cp\u003eType\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c3\"\u003e \u003cp\u003eSpecies\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c4\"\u003e \u003cp\u003eNumber\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c5\"\u003e \u003cp\u003eDBH\u0026thinsp;\u0026plusmn;\u0026thinsp;sd(cm)\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c6\"\u003e \u003cp\u003eTH\u0026thinsp;\u0026plusmn;\u0026thinsp;sd(m)\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c7\"\u003e \u003cp\u003eCW\u0026thinsp;\u0026plusmn;\u0026thinsp;sd(m)\u003c/p\u003e \u003c/th\u003e \u003c/tr\u003e \u003c/thead\u003e \u003ctbody\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\" morerows=\"8\" rowspan=\"9\"\u003e \u003cp\u003e\u003cb\u003eML stand\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cb\u003eplanted trees\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e\u003cem\u003eML\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e488\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c5\"\u003e \u003cp\u003e22.39\u0026thinsp;\u0026plusmn;\u0026thinsp;6.7\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c6\"\u003e \u003cp\u003e20.36\u0026thinsp;\u0026plusmn;\u0026thinsp;4.8\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c7\"\u003e \u003cp\u003e3.95\u0026thinsp;\u0026plusmn;\u0026thinsp;2.05\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c2\" morerows=\"3\" rowspan=\"4\"\u003e \u003cp\u003e\u003cb\u003ereg.(shade-tolerant)\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e\u003cem\u003eAP\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e26\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c5\"\u003e \u003cp\u003e0.80\u0026thinsp;\u0026plusmn;\u0026thinsp;0.44\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c6\"\u003e \u003cp\u003e0.91\u0026thinsp;\u0026plusmn;\u0026thinsp;0.55\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c7\"\u003e \u003cp\u003e0.25\u0026thinsp;\u0026plusmn;\u0026thinsp;0.18\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e\u003cem\u003eML\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e145\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c5\"\u003e \u003cp\u003e3.00\u0026thinsp;\u0026plusmn;\u0026thinsp;3.12\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c6\"\u003e \u003cp\u003e4.08\u0026thinsp;\u0026plusmn;\u0026thinsp;3.44\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c7\"\u003e \u003cp\u003e0.89\u0026thinsp;\u0026plusmn;\u0026thinsp;0.86\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e\u003cem\u003eMM-Da\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e23\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c5\"\u003e \u003cp\u003e1.04\u0026thinsp;\u0026plusmn;\u0026thinsp;0.78\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c6\"\u003e \u003cp\u003e1.13\u0026thinsp;\u0026plusmn;\u0026thinsp;0.77\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c7\"\u003e \u003cp\u003e0.29\u0026thinsp;\u0026plusmn;\u0026thinsp;0.3\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e\u003cem\u003eOthers\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e25\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c5\"\u003e \u003cp\u003e1.22\u0026thinsp;\u0026plusmn;\u0026thinsp;0.69\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c6\"\u003e \u003cp\u003e1.26\u0026thinsp;\u0026plusmn;\u0026thinsp;0.76\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c7\"\u003e \u003cp\u003e0.31\u0026thinsp;\u0026plusmn;\u0026thinsp;0.25\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c2\" morerows=\"3\" rowspan=\"4\"\u003e \u003cp\u003e\u003cb\u003ereg.(shade-intolerant)\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e\u003cem\u003eAP\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e25\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c5\"\u003e \u003cp\u003e0.58\u0026thinsp;\u0026plusmn;\u0026thinsp;0.20\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c6\"\u003e \u003cp\u003e0.86\u0026thinsp;\u0026plusmn;\u0026thinsp;0.39\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c7\"\u003e \u003cp\u003e0.2\u0026thinsp;\u0026plusmn;\u0026thinsp;0.09\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e\u003cb\u003eML\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e98\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c5\"\u003e \u003cp\u003e2.47\u0026thinsp;\u0026plusmn;\u0026thinsp;2.63\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c6\"\u003e \u003cp\u003e3.90\u0026thinsp;\u0026plusmn;\u0026thinsp;2.79\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c7\"\u003e \u003cp\u003e0.83\u0026thinsp;\u0026plusmn;\u0026thinsp;0.75\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e\u003cb\u003eMM-Da\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e32\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c5\"\u003e \u003cp\u003e1.10\u0026thinsp;\u0026plusmn;\u0026thinsp;0.82\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c6\"\u003e \u003cp\u003e1.05\u0026thinsp;\u0026plusmn;\u0026thinsp;0.71\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c7\"\u003e \u003cp\u003e0.3\u0026thinsp;\u0026plusmn;\u0026thinsp;0.26\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e\u003cem\u003eOthers\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e27\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c5\"\u003e \u003cp\u003e1.12\u0026thinsp;\u0026plusmn;\u0026thinsp;0.85\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c6\"\u003e \u003cp\u003e1.25\u0026thinsp;\u0026plusmn;\u0026thinsp;0.99\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c7\"\u003e \u003cp\u003e0.4\u0026thinsp;\u0026plusmn;\u0026thinsp;0.38\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\" morerows=\"23\" rowspan=\"24\"\u003e \u003cp\u003e\u003cb\u003eMM-Da stand\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cb\u003eplanted trees\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e\u003cem\u003eMM-Da\u003c/em\u003e\u003c/p\u003e \u003cp\u003e\u003cem\u003eHH\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e206\u003c/p\u003e \u003cp\u003e2\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c5\"\u003e \u003cp\u003e21.42\u0026thinsp;\u0026plusmn;\u0026thinsp;7.7\u003c/p\u003e \u003cp\u003e36.91\u0026thinsp;\u0026plusmn;\u0026thinsp;24.8\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c6\"\u003e \u003cp\u003e19.92\u0026thinsp;\u0026plusmn;\u0026thinsp;4.3\u003c/p\u003e \u003cp\u003e16.0\u0026thinsp;\u0026plusmn;\u0026thinsp;5.6\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c7\"\u003e \u003cp\u003e4.29\u0026thinsp;\u0026plusmn;\u0026thinsp;1.96\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c2\" morerows=\"10\" rowspan=\"11\"\u003e \u003cp\u003e\u003cb\u003ereg.(shade-tolerant)\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e\u003cem\u003eAP\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e42\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c5\"\u003e \u003cp\u003e1.31\u0026thinsp;\u0026plusmn;\u0026thinsp;0.88\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c6\"\u003e \u003cp\u003e1.60\u0026thinsp;\u0026plusmn;\u0026thinsp;1.13\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c7\"\u003e \u003cp\u003e0.37\u0026thinsp;\u0026plusmn;\u0026thinsp;0.21\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e\u003cem\u003eAQ\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e54\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c5\"\u003e \u003cp\u003e0.99\u0026thinsp;\u0026plusmn;\u0026thinsp;0.43\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c6\"\u003e \u003cp\u003e1.47\u0026thinsp;\u0026plusmn;\u0026thinsp;0.82\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c7\"\u003e \u003cp\u003e0.46\u0026thinsp;\u0026plusmn;\u0026thinsp;0.26\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e\u003cem\u003eDC\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e58\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c5\"\u003e \u003cp\u003e0.60\u0026thinsp;\u0026plusmn;\u0026thinsp;0.31\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c6\"\u003e \u003cp\u003e1.00\u0026thinsp;\u0026plusmn;\u0026thinsp;0.35\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c7\"\u003e \u003cp\u003e0.43\u0026thinsp;\u0026plusmn;\u0026thinsp;0.17\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e\u003cem\u003eFH\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e12\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c5\"\u003e \u003cp\u003e0.69\u0026thinsp;\u0026plusmn;\u0026thinsp;0.28\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c6\"\u003e \u003cp\u003e1.37\u0026thinsp;\u0026plusmn;\u0026thinsp;0.55\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c7\"\u003e \u003cp\u003e0.23\u0026thinsp;\u0026plusmn;\u0026thinsp;0.17\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e\u003cb\u003eML\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e47\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c5\"\u003e \u003cp\u003e1.75\u0026thinsp;\u0026plusmn;\u0026thinsp;0.91\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c6\"\u003e \u003cp\u003e2.96\u0026thinsp;\u0026plusmn;\u0026thinsp;1.72\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c7\"\u003e \u003cp\u003e0.82\u0026thinsp;\u0026plusmn;\u0026thinsp;0.65\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e\u003cb\u003eMM-Da\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e139\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c5\"\u003e \u003cp\u003e1.29\u0026thinsp;\u0026plusmn;\u0026thinsp;1.01\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c6\"\u003e \u003cp\u003e1.95\u0026thinsp;\u0026plusmn;\u0026thinsp;1.49\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c7\"\u003e \u003cp\u003e0.55\u0026thinsp;\u0026plusmn;\u0026thinsp;0.46\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e\u003cem\u003eMP-H\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e35\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c5\"\u003e \u003cp\u003e0.83\u0026thinsp;\u0026plusmn;\u0026thinsp;0.59\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c6\"\u003e \u003cp\u003e1.31\u0026thinsp;\u0026plusmn;\u0026thinsp;0.67\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c7\"\u003e \u003cp\u003e0.3\u0026thinsp;\u0026plusmn;\u0026thinsp;0.17\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e\u003cem\u003eOP\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e10\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c5\"\u003e \u003cp\u003e1.16\u0026thinsp;\u0026plusmn;\u0026thinsp;0.12\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c6\"\u003e \u003cp\u003e1.15\u0026thinsp;\u0026plusmn;\u0026thinsp;0.54\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c7\"\u003e \u003cp\u003e0.31\u0026thinsp;\u0026plusmn;\u0026thinsp;0.24\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e\u003cem\u003ePR\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e17\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c5\"\u003e \u003cp\u003e0.81\u0026thinsp;\u0026plusmn;\u0026thinsp;0.47\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c6\"\u003e \u003cp\u003e1.23\u0026thinsp;\u0026plusmn;\u0026thinsp;0.59\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c7\"\u003e \u003cp\u003e0.27\u0026thinsp;\u0026plusmn;\u0026thinsp;0.17\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e\u003cem\u003eSG-R\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e11\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c5\"\u003e \u003cp\u003e1.49\u0026thinsp;\u0026plusmn;\u0026thinsp;0.23\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c6\"\u003e \u003cp\u003e1.17\u0026thinsp;\u0026plusmn;\u0026thinsp;0.58\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c7\"\u003e \u003cp\u003e0.32\u0026thinsp;\u0026plusmn;\u0026thinsp;0.15\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e\u003cem\u003eOther\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e75\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c5\"\u003e \u003cp\u003e1.26\u0026thinsp;\u0026plusmn;\u0026thinsp;0.83\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c6\"\u003e \u003cp\u003e1.73\u0026thinsp;\u0026plusmn;\u0026thinsp;1.03\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c7\"\u003e \u003cp\u003e0.64\u0026thinsp;\u0026plusmn;\u0026thinsp;0.85\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c2\" morerows=\"11\" rowspan=\"12\"\u003e \u003cp\u003e\u003cb\u003ereg.(shade-intolerant)\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e\u003cem\u003eAP\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e66\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c5\"\u003e \u003cp\u003e0.81\u0026thinsp;\u0026plusmn;\u0026thinsp;0.32\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c6\"\u003e \u003cp\u003e1.11\u0026thinsp;\u0026plusmn;\u0026thinsp;0.39\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c7\"\u003e \u003cp\u003e0.29\u0026thinsp;\u0026plusmn;\u0026thinsp;0.15\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e\u003cem\u003eAQ\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e33\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c5\"\u003e \u003cp\u003e1.17\u0026thinsp;\u0026plusmn;\u0026thinsp;0.70\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c6\"\u003e \u003cp\u003e1.58\u0026thinsp;\u0026plusmn;\u0026thinsp;0.81\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c7\"\u003e \u003cp\u003e0.48\u0026thinsp;\u0026plusmn;\u0026thinsp;0.3\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e\u003cem\u003eDC\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e121\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c5\"\u003e \u003cp\u003e0.72\u0026thinsp;\u0026plusmn;\u0026thinsp;0.24\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c6\"\u003e \u003cp\u003e0.96\u0026thinsp;\u0026plusmn;\u0026thinsp;0.38\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c7\"\u003e \u003cp\u003e0.4\u0026thinsp;\u0026plusmn;\u0026thinsp;0.2\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e\u003cem\u003eFH\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e14\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c5\"\u003e \u003cp\u003e0.54\u0026thinsp;\u0026plusmn;\u0026thinsp;0.20\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c6\"\u003e \u003cp\u003e1.03\u0026thinsp;\u0026plusmn;\u0026thinsp;0.41\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c7\"\u003e \u003cp\u003e0.16\u0026thinsp;\u0026plusmn;\u0026thinsp;0.1\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e\u003cem\u003eHH\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e10\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c5\"\u003e \u003cp\u003e2.13\u0026thinsp;\u0026plusmn;\u0026thinsp;0.77\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c6\"\u003e \u003cp\u003e1.99\u0026thinsp;\u0026plusmn;\u0026thinsp;1.22\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c7\"\u003e \u003cp\u003e0.49\u0026thinsp;\u0026plusmn;\u0026thinsp;0.31\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e\u003cb\u003eML\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e14\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c5\"\u003e \u003cp\u003e1.92\u0026thinsp;\u0026plusmn;\u0026thinsp;1.22\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c6\"\u003e \u003cp\u003e3.35\u0026thinsp;\u0026plusmn;\u0026thinsp;1.40\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c7\"\u003e \u003cp\u003e1.11\u0026thinsp;\u0026plusmn;\u0026thinsp;0.8\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e\u003cb\u003eMM-Da\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e164\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c5\"\u003e \u003cp\u003e1.42\u0026thinsp;\u0026plusmn;\u0026thinsp;0.91\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c6\"\u003e \u003cp\u003e1.68\u0026thinsp;\u0026plusmn;\u0026thinsp;1.06\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c7\"\u003e \u003cp\u003e0.52\u0026thinsp;\u0026plusmn;\u0026thinsp;0.34\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e\u003cem\u003eMP-H\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e72\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c5\"\u003e \u003cp\u003e0.83\u0026thinsp;\u0026plusmn;\u0026thinsp;0.82\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c6\"\u003e \u003cp\u003e1.08\u0026thinsp;\u0026plusmn;\u0026thinsp;0.38\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c7\"\u003e \u003cp\u003e0.24\u0026thinsp;\u0026plusmn;\u0026thinsp;0.15\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e\u003cem\u003ePR\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e17\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c5\"\u003e \u003cp\u003e0.58\u0026thinsp;\u0026plusmn;\u0026thinsp;0.13\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c6\"\u003e \u003cp\u003e1.16\u0026thinsp;\u0026plusmn;\u0026thinsp;0.38\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c7\"\u003e \u003cp\u003e0.31\u0026thinsp;\u0026plusmn;\u0026thinsp;0.19\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e\u003cem\u003eSG-R\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e16\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c5\"\u003e \u003cp\u003e0.56\u0026thinsp;\u0026plusmn;\u0026thinsp;0.04\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c6\"\u003e \u003cp\u003e0.94\u0026thinsp;\u0026plusmn;\u0026thinsp;0.29\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c7\"\u003e \u003cp\u003e0.36\u0026thinsp;\u0026plusmn;\u0026thinsp;0.24\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e\u003cem\u003eWU\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e12\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c5\"\u003e \u003cp\u003e1.76\u0026thinsp;\u0026plusmn;\u0026thinsp;1.10\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c6\"\u003e \u003cp\u003e2.40\u0026thinsp;\u0026plusmn;\u0026thinsp;1.54\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c7\"\u003e \u003cp\u003e0.8\u0026thinsp;\u0026plusmn;\u0026thinsp;0.61\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e\u003cem\u003eOthers\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e71\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c5\"\u003e \u003cp\u003e1.04\u0026thinsp;\u0026plusmn;\u0026thinsp;0.87\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c6\"\u003e \u003cp\u003e1.51\u0026thinsp;\u0026plusmn;\u0026thinsp;0.85\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c7\"\u003e \u003cp\u003e0.4\u0026thinsp;\u0026plusmn;\u0026thinsp;0.33\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\" morerows=\"19\" rowspan=\"20\"\u003e \u003cp\u003e\u003cb\u003eMixed\u003c/b\u003e\u003c/p\u003e \u003cp\u003e\u003cb\u003estand\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\" morerows=\"1\" rowspan=\"2\"\u003e \u003cp\u003e\u003cb\u003eplanted trees\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e\u003cem\u003eML\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e59\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c5\"\u003e \u003cp\u003e21.51\u0026thinsp;\u0026plusmn;\u0026thinsp;6.7\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c6\"\u003e \u003cp\u003e22.88\u0026thinsp;\u0026plusmn;\u0026thinsp;5.1\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c7\"\u003e \u003cp\u003e3.64\u0026thinsp;\u0026plusmn;\u0026thinsp;1.06\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e\u003cem\u003eMM-Da\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e141\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c5\"\u003e \u003cp\u003e22.60\u0026thinsp;\u0026plusmn;\u0026thinsp;8.0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c6\"\u003e \u003cp\u003e21.08\u0026thinsp;\u0026plusmn;\u0026thinsp;3.7\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c7\"\u003e \u003cp\u003e3.84\u0026thinsp;\u0026plusmn;\u0026thinsp;1.31\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c2\" morerows=\"9\" rowspan=\"10\"\u003e \u003cp\u003e\u003cb\u003ereg.(shade-tolerant)\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e\u003cem\u003eAP\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e36\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c5\"\u003e \u003cp\u003e0.66\u0026thinsp;\u0026plusmn;\u0026thinsp;0.29\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c6\"\u003e \u003cp\u003e1.28\u0026thinsp;\u0026plusmn;\u0026thinsp;1.76\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c7\"\u003e \u003cp\u003e0.31\u0026thinsp;\u0026plusmn;\u0026thinsp;0.12\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e\u003cem\u003eDC\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e24\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c5\"\u003e \u003cp\u003e0.83\u0026thinsp;\u0026plusmn;\u0026thinsp;0.56\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c6\"\u003e \u003cp\u003e1.01\u0026thinsp;\u0026plusmn;\u0026thinsp;0.6\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c7\"\u003e \u003cp\u003e0.29\u0026thinsp;\u0026plusmn;\u0026thinsp;0.18\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e\u003cem\u003eER\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e30\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c5\"\u003e \u003cp\u003e0.57\u0026thinsp;\u0026plusmn;\u0026thinsp;0.27\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c6\"\u003e \u003cp\u003e1.03\u0026thinsp;\u0026plusmn;\u0026thinsp;0.52\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c7\"\u003e \u003cp\u003e0.22\u0026thinsp;\u0026plusmn;\u0026thinsp;0.13\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e\u003cem\u003eLN\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e11\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c5\"\u003e \u003cp\u003e0.58\u0026thinsp;\u0026plusmn;\u0026thinsp;0.29\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c6\"\u003e \u003cp\u003e1.09\u0026thinsp;\u0026plusmn;\u0026thinsp;0.5\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c7\"\u003e \u003cp\u003e0.33\u0026thinsp;\u0026plusmn;\u0026thinsp;0.15\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e\u003cem\u003eMJ-J\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e12\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c5\"\u003e \u003cp\u003e2.11\u0026thinsp;\u0026plusmn;\u0026thinsp;1.25\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c6\"\u003e \u003cp\u003e1.82\u0026thinsp;\u0026plusmn;\u0026thinsp;1.61\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c7\"\u003e \u003cp\u003e0.55\u0026thinsp;\u0026plusmn;\u0026thinsp;0.45\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e\u003cb\u003eML\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e367\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c5\"\u003e \u003cp\u003e1.51\u0026thinsp;\u0026plusmn;\u0026thinsp;1.00\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c6\"\u003e \u003cp\u003e2.34\u0026thinsp;\u0026plusmn;\u0026thinsp;1.32\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c7\"\u003e \u003cp\u003e0.66\u0026thinsp;\u0026plusmn;\u0026thinsp;0.54\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e\u003cb\u003eMM-Da\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e53\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c5\"\u003e \u003cp\u003e2.76\u0026thinsp;\u0026plusmn;\u0026thinsp;3.40\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c6\"\u003e \u003cp\u003e2.01\u0026thinsp;\u0026plusmn;\u0026thinsp;2.51\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c7\"\u003e \u003cp\u003e0.66\u0026thinsp;\u0026plusmn;\u0026thinsp;0.79\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e\u003cem\u003eMP-H\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e17\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c5\"\u003e \u003cp\u003e0.74\u0026thinsp;\u0026plusmn;\u0026thinsp;0.43\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c6\"\u003e \u003cp\u003e0.85\u0026thinsp;\u0026plusmn;\u0026thinsp;0.52\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c7\"\u003e \u003cp\u003e0.26\u0026thinsp;\u0026plusmn;\u0026thinsp;0.17\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e\u003cem\u003eWZⅨ\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e12\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c5\"\u003e \u003cp\u003e0.74\u0026thinsp;\u0026plusmn;\u0026thinsp;0.40\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c6\"\u003e \u003cp\u003e0.97\u0026thinsp;\u0026plusmn;\u0026thinsp;0.56\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c7\"\u003e \u003cp\u003e0.36\u0026thinsp;\u0026plusmn;\u0026thinsp;0.31\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e\u003cem\u003eOther\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e105\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c5\"\u003e \u003cp\u003e1.48\u0026thinsp;\u0026plusmn;\u0026thinsp;1.0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c6\"\u003e \u003cp\u003e1.25\u0026thinsp;\u0026plusmn;\u0026thinsp;0.9\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c7\"\u003e \u003cp\u003e0.39\u0026thinsp;\u0026plusmn;\u0026thinsp;0.43\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c2\" morerows=\"7\" rowspan=\"8\"\u003e \u003cp\u003e\u003cb\u003ereg.(shade-intolerant)\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e\u003cem\u003eAP\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e29\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c5\"\u003e \u003cp\u003e0.88\u0026thinsp;\u0026plusmn;\u0026thinsp;0.54\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c6\"\u003e \u003cp\u003e1.31\u0026thinsp;\u0026plusmn;\u0026thinsp;0.65\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c7\"\u003e \u003cp\u003e0.37\u0026thinsp;\u0026plusmn;\u0026thinsp;0.22\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e\u003cem\u003eDC\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e14\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c5\"\u003e \u003cp\u003e0.74\u0026thinsp;\u0026plusmn;\u0026thinsp;0.58\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c6\"\u003e \u003cp\u003e0.82\u0026thinsp;\u0026plusmn;\u0026thinsp;0.52\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c7\"\u003e \u003cp\u003e0.26\u0026thinsp;\u0026plusmn;\u0026thinsp;0.1\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e\u003cem\u003eLN\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e12\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c5\"\u003e \u003cp\u003e0.61\u0026thinsp;\u0026plusmn;\u0026thinsp;0.42\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c6\"\u003e \u003cp\u003e1.17\u0026thinsp;\u0026plusmn;\u0026thinsp;0.29\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c7\"\u003e \u003cp\u003e0.25\u0026thinsp;\u0026plusmn;\u0026thinsp;0.09\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e\u003cb\u003eML\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e305\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c5\"\u003e \u003cp\u003e1.42\u0026thinsp;\u0026plusmn;\u0026thinsp;0.84\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c6\"\u003e \u003cp\u003e2.21\u0026thinsp;\u0026plusmn;\u0026thinsp;1.23\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c7\"\u003e \u003cp\u003e0.61\u0026thinsp;\u0026plusmn;\u0026thinsp;0.46\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e\u003cb\u003eMM-Da\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e47\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c5\"\u003e \u003cp\u003e1.85\u0026thinsp;\u0026plusmn;\u0026thinsp;2.77\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c6\"\u003e \u003cp\u003e1.52\u0026thinsp;\u0026plusmn;\u0026thinsp;1.62\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c7\"\u003e \u003cp\u003e0.45\u0026thinsp;\u0026plusmn;\u0026thinsp;0.45\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e\u003cem\u003eMP-H\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e20\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c5\"\u003e \u003cp\u003e0.48\u0026thinsp;\u0026plusmn;\u0026thinsp;0.05\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c6\"\u003e \u003cp\u003e0.80\u0026thinsp;\u0026plusmn;\u0026thinsp;0.33\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c7\"\u003e \u003cp\u003e0.25\u0026thinsp;\u0026plusmn;\u0026thinsp;0.13\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e\u003cem\u003ePR\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e23\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c5\"\u003e \u003cp\u003e0.63\u0026thinsp;\u0026plusmn;\u0026thinsp;0.33\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c6\"\u003e \u003cp\u003e1.27\u0026thinsp;\u0026plusmn;\u0026thinsp;0.45\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c7\"\u003e \u003cp\u003e0.4\u0026thinsp;\u0026plusmn;\u0026thinsp;0.16\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e\u003cem\u003eOthers\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e79\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c5\"\u003e \u003cp\u003e1.61\u0026thinsp;\u0026plusmn;\u0026thinsp;1.20\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c6\"\u003e \u003cp\u003e1.21\u0026thinsp;\u0026plusmn;\u0026thinsp;0.92\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c7\"\u003e \u003cp\u003e0.4\u0026thinsp;\u0026plusmn;\u0026thinsp;0.38\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003c/tbody\u003e \u003c/colgroup\u003e \u003c/table\u003e\u003c/div\u003e \u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec7\" class=\"Section3\"\u003e \u003ch2\u003e2.3.2 Point Pattern Analysis\u003c/h2\u003e \u003cp\u003eThe pair correlation function which is based on the distance between a pair of points (PCF, \u003cspan class=\"InlineEquation\"\u003e\u003cspan class=\"mathinline\"\u003e\\(\\:g\\left(r\\right)=\\frac{\\widehat{K}\\left(r\\right)}{2\\pi\\:r}\\)\u003c/span\u003e\u003c/span\u003e, \u003cspan class=\"InlineEquation\"\u003e\u003cspan class=\"mathinline\"\u003e\\(\\:\\widehat{K}\\left(r\\right)=A\\sum\\:_{i=1}^{N}\\sum\\:_{j=1}^{N}\\frac{{w}_{ij}\\left(d\\right)}{{N}^{2}}\\)\u003c/span\u003e\u003c/span\u003e, \u003cem\u003eA\u003c/em\u003e\u0026thinsp;=\u0026thinsp;area; \u003cem\u003eN\u003c/em\u003e\u0026thinsp;=\u0026thinsp;number of trees; \u003cem\u003er\u003c/em\u003e\u0026thinsp;=\u0026thinsp;the observation scale (0, 1, 2\u0026hellip; 20 m); λ\u0026thinsp;=\u0026thinsp;density; \u003cem\u003ew\u003c/em\u003e\u003csub\u003e\u003cem\u003eij\u003c/em\u003e\u003c/sub\u003e is the weight.) can analyze tree spatial distribution patterns across scales and to identify the scales at which patterns may change, thereby facilitating the interpretation of spatial patterns (Getzin et al., \u003cspan citationid=\"CR35\" class=\"CitationRef\"\u003e2006\u003c/span\u003e; Wiegand and Moloney, \u003cspan citationid=\"CR127\" class=\"CitationRef\"\u003e2014\u003c/span\u003e). Therefore, we used the univariate distribution model \u003cem\u003eg\u003c/em\u003e\u003csub\u003e11\u003c/sub\u003e(\u003cem\u003er\u003c/em\u003e) of PCF to analyze the spatial distributions of stands, planted trees, and regenerations. We adopted complete spatial randomness (CSR) as the null model (Wiegand and Moloney, \u003cspan citationid=\"CR127\" class=\"CitationRef\"\u003e2014\u003c/span\u003e), and the Monte Carlo (MC) method was used to simulate the observed values 199 times. The observation scale was set to half the short side of the quadrat (\u003cem\u003er\u003c/em\u003e\u0026thinsp;=\u0026thinsp;0\u0026ndash;20 m) and the envelope curves were plotted using 5% of the maximum and minimum values. Observations falling above, within, and below the simulation interval indicate significant aggregation, randomness, and regularity of the distribution pattern at the corresponding scales, respectively (Li et al., \u003cspan citationid=\"CR57\" class=\"CitationRef\"\u003e2021b\u003c/span\u003e).\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec8\" class=\"Section3\"\u003e \u003ch2\u003e2.3.3 Analysis of Spatial Correlation\u003c/h2\u003e \u003cp\u003ePlantations have a clear origin, with unified planting time and locations, high management intensity, and relatively homogeneous habitats (Bremer and Farley, \u003cspan citationid=\"CR11\" class=\"CitationRef\"\u003e2010\u003c/span\u003e; Paquette and Messier, \u003cspan citationid=\"CR85\" class=\"CitationRef\"\u003e2010\u003c/span\u003e), even small plots can reveal interspecific spatial correlations. Therefore, we applied the case-control method to analyze the spatial correlation between planted trees and regenerations (Wiegand and Moloney, \u003cspan citationid=\"CR123\" class=\"CitationRef\"\u003e2004\u003c/span\u003e; Diggle et al., \u003cspan citationid=\"CR29\" class=\"CitationRef\"\u003e2007\u003c/span\u003e), the coordinates of planted trees were fixed as the control (Pattern 1), whereas regenerations were treated as the case (Pattern 2) and were randomly distributed around the control, the results will reflect the influence of planted trees on the distribution of regenerations. We used the bivariate distribution model \u003cem\u003eg\u003c/em\u003e\u003csub\u003e12\u003c/sub\u003e(\u003cem\u003er\u003c/em\u003e) to perform this analysis. In addition, using CSR as the null model, we treated the shade-tolerant group as Pattern 1 and the shade-intolerant group as Pattern 2, and analyzed spatial correlations between functional groups using \u003cem\u003eg\u003c/em\u003e\u003csub\u003e12\u003c/sub\u003e(\u003cem\u003er\u003c/em\u003e) and \u003cem\u003eg\u003c/em\u003e\u003csub\u003e21\u003c/sub\u003e(\u003cem\u003er\u003c/em\u003e), the parameter settings were consistent with those for the univariate distribution model \u003cem\u003eg\u003c/em\u003e\u003csub\u003e11\u003c/sub\u003e(\u003cem\u003er\u003c/em\u003e). At the observation scale \u003cem\u003er\u003c/em\u003e, \u003cem\u003eg\u003c/em\u003e\u003csub\u003e12\u003c/sub\u003e(\u003cem\u003er\u003c/em\u003e) represents the distribution of Pattern 2 around Pattern 1, i.e., the spatial correlation of Pattern 2 relative to Pattern 1. Similarly, at the observation scale \u003cem\u003er\u003c/em\u003e, \u003cem\u003eg\u003c/em\u003e\u003csub\u003e21\u003c/sub\u003e(\u003cem\u003er\u003c/em\u003e) represents the spatial correlation of Pattern 1 relative to Pattern 2. The theoretical standard value for spatial correlation is 1, when \u003cem\u003eg\u003c/em\u003e\u003csub\u003e12\u003c/sub\u003e(\u003cem\u003er\u003c/em\u003e) and \u003cem\u003eg\u003c/em\u003e\u003csub\u003e21\u003c/sub\u003e(\u003cem\u003er\u003c/em\u003e)\u0026thinsp;\u0026gt;\u0026thinsp;1, the correlation is positive, that means attraction, when \u003cem\u003eg\u003c/em\u003e\u003csub\u003e12\u003c/sub\u003e(\u003cem\u003er\u003c/em\u003e) and \u003cem\u003eg\u003c/em\u003e\u003csub\u003e21\u003c/sub\u003e(\u003cem\u003er\u003c/em\u003e)\u0026thinsp;\u0026lt;\u0026thinsp;1, the correlation is negative, that means repulsion, and when \u003cem\u003eg\u003c/em\u003e\u003csub\u003e12\u003c/sub\u003e(\u003cem\u003er\u003c/em\u003e) and \u003cem\u003eg\u003c/em\u003e\u003csub\u003e21\u003c/sub\u003e(\u003cem\u003er\u003c/em\u003e)\u0026thinsp;=\u0026thinsp;1, there is no correlation, that means independence (Getzin et al., \u003cspan citationid=\"CR35\" class=\"CitationRef\"\u003e2006\u003c/span\u003e; Wiegand and Moloney, \u003cspan citationid=\"CR127\" class=\"CitationRef\"\u003e2014\u003c/span\u003e). When the observed values exceeded the 95% simulation envelopes, the spatial correlation is considered significant, otherwise the relationship is regarded as weak. All analyses were conducted in Programita (version, 2018) (Wiegand and Moloney, \u003cspan citationid=\"CR127\" class=\"CitationRef\"\u003e2014\u003c/span\u003e), and graphing were produced in R (version, 4.4.3) using the \u003cem\u003eggplot2\u003c/em\u003e package.\u003c/p\u003e \u003c/div\u003e \u003c/div\u003e"},{"header":"3. Results","content":"\u003cdiv id=\"Sec10\" class=\"Section2\"\u003e \u003ch2\u003e3.1 Spatial Distribution Patterns of \u003cem\u003eM. macclurei\u003c/em\u003e and \u003cem\u003eM. laosensis\u003c/em\u003e Plantations\u003c/h2\u003e \u003cp\u003eIn both monoculture stands and the mixed stand showed aggregated distribution patterns, but the scales differed (Fig.\u0026nbsp;\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e2\u003c/span\u003ea-\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e2\u003c/span\u003ec). Among them, the MM-Da Stand was aggregated at \u003cem\u003er\u003c/em\u003e\u0026thinsp;=\u0026thinsp;0\u0026ndash;16 m, whereas the ML Stand and the Mixed Stand were aggregated at \u003cem\u003er\u003c/em\u003e\u0026thinsp;=\u0026thinsp;0\u0026ndash;12 m. In the ML Stand, planted trees were aggregated at \u003cem\u003er\u003c/em\u003e\u0026thinsp;=\u0026thinsp;0\u0026ndash;2 m and were randomly distributed at other scales (Fig.\u0026nbsp;\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e2\u003c/span\u003ed). In the MM-Da Stand and the Mixed Stand, planted trees were aggregated at \u003cem\u003er\u003c/em\u003e\u0026thinsp;=\u0026thinsp;0\u0026ndash;3 m and \u003cem\u003er\u003c/em\u003e\u0026thinsp;=\u0026thinsp;0\u0026ndash;2 m, respectively (Fig.\u0026nbsp;\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e2\u003c/span\u003ee-\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e2\u003c/span\u003ef). Regenerations in the ML Stand was aggregated at \u003cem\u003er\u003c/em\u003e\u0026thinsp;=\u0026thinsp;0\u0026ndash;14 m (Fig.\u0026nbsp;\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e2\u003c/span\u003eg), whereas regenerations in the MM-Da Stand was aggregated at almost all scales (Fig.\u0026nbsp;\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e2\u003c/span\u003eh). In the Mixed Stand, the aggregation of regenerations was similar to that in the ML Stand (Fig.\u0026nbsp;\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e2\u003c/span\u003ei).\u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec11\" class=\"Section2\"\u003e \u003ch2\u003e3.2 Spatial Distribution Patterns of Functional groups\u003c/h2\u003e \u003cp\u003eIn both monoculture stands, the shade-tolerant group was aggregated at all observation scales (Fig.\u0026nbsp;\u003cspan refid=\"Fig3\" class=\"InternalRef\"\u003e3\u003c/span\u003ea-\u003cspan refid=\"Fig3\" class=\"InternalRef\"\u003e3\u003c/span\u003eb). In contrast, in the Mixed Stand, it was aggregated only at \u003cem\u003er\u003c/em\u003e\u0026thinsp;=\u0026thinsp;0\u0026ndash;8 m (Fig.\u0026nbsp;\u003cspan refid=\"Fig3\" class=\"InternalRef\"\u003e3\u003c/span\u003ec). In the ML Stand, the shade-intolerant group was mainly aggregated at \u003cem\u003er\u003c/em\u003e\u0026thinsp;=\u0026thinsp;0\u0026ndash;13 m (Fig.\u0026nbsp;\u003cspan refid=\"Fig3\" class=\"InternalRef\"\u003e3\u003c/span\u003ed). However, in the MM-Da Stand, its aggregation scale expanded to \u003cem\u003er\u003c/em\u003e\u0026thinsp;=\u0026thinsp;0\u0026ndash;20 m (Fig.\u0026nbsp;\u003cspan refid=\"Fig3\" class=\"InternalRef\"\u003e3\u003c/span\u003ee). Moreover, the aggregation intensity of the shade-intolerant group was lower in the Mixed Stand than in monoculture stands, occurring only in the range of \u003cem\u003er\u003c/em\u003e\u0026thinsp;=\u0026thinsp;0\u0026ndash;7 m, while exhibiting a random distribution in the range of \u003cem\u003er\u003c/em\u003e\u0026thinsp;=\u0026thinsp;8\u0026ndash;16 m, and regular at the remaining scales (Fig.\u0026nbsp;\u003cspan refid=\"Fig3\" class=\"InternalRef\"\u003e3\u003c/span\u003ef).\u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec12\" class=\"Section2\"\u003e \u003ch2\u003e3.3 Spatial Correlations between Planted trees and Regeneration\u003c/h2\u003e \u003cp\u003eIn the ML Stand and the Mixed Stand, planted trees and regeneration were positively associated at \u003cem\u003er\u003c/em\u003e\u0026thinsp;=\u0026thinsp;0\u0026ndash;2 m, but in the MM-Da Stand, they were positively associated at \u003cem\u003er\u003c/em\u003e\u0026thinsp;=\u0026thinsp;0\u0026ndash;2 m but negatively associated at other scales (Fig.\u0026nbsp;\u003cspan refid=\"Fig4\" class=\"InternalRef\"\u003e4\u003c/span\u003ea-\u003cspan refid=\"Fig4\" class=\"InternalRef\"\u003e4\u003c/span\u003ec). Under both planting patterns, planted trees and the shade-tolerant group were positively associated at small scales (\u003cem\u003er\u003c/em\u003e\u0026thinsp;=\u0026thinsp;0\u0026ndash;5 m), whereas they showed random correlation at other scales (Fig.\u0026nbsp;\u003cspan refid=\"Fig4\" class=\"InternalRef\"\u003e4\u003c/span\u003ed-\u003cspan refid=\"Fig4\" class=\"InternalRef\"\u003e4\u003c/span\u003ef). In the ML Stand, planted trees and the shade-intolerant group were repulsive at \u003cem\u003er\u003c/em\u003e\u0026thinsp;=\u0026thinsp;0\u0026ndash;3 m, and they were independent at the remaining scales (Fig.\u0026nbsp;\u003cspan refid=\"Fig4\" class=\"InternalRef\"\u003e4\u003c/span\u003eg). The spatial correlation between planted trees and the shade-intolerant group was negative in the MM-Da Stand (Fig.\u0026nbsp;\u003cspan refid=\"Fig4\" class=\"InternalRef\"\u003e4\u003c/span\u003eh), but in the Mixed Stand, it was negative at \u003cem\u003er\u003c/em\u003e\u0026thinsp;=\u0026thinsp;0\u0026ndash;4 m but was mainly random at the remaining scales (Fig.\u0026nbsp;\u003cspan refid=\"Fig4\" class=\"InternalRef\"\u003e4\u003c/span\u003ei).\u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec13\" class=\"Section2\"\u003e \u003ch2\u003e3.4 Spatial Correlations between Functional Groups\u003c/h2\u003e \u003cp\u003eThe spatial correlation between the shade-tolerant and shade-intolerant groups was negative at small scales in all stands, but their relationships at other scales differed slightly (Fig.\u0026nbsp;\u003cspan refid=\"Fig5\" class=\"InternalRef\"\u003e5\u003c/span\u003ea-\u003cspan refid=\"Fig5\" class=\"InternalRef\"\u003e5\u003c/span\u003ec). Specifically, the two groups showed attraction at \u003cem\u003er\u003c/em\u003e\u0026thinsp;=\u0026thinsp;4\u0026ndash;20 m and \u003cem\u003er\u003c/em\u003e\u0026thinsp;=\u0026thinsp;6\u0026ndash;20 m in both monoculture stands, respectively. In contrast, in the Mixed Stand, they showed attraction at \u003cem\u003er\u003c/em\u003e\u0026thinsp;=\u0026thinsp;4\u0026ndash;10 m but repulsion at \u003cem\u003er\u003c/em\u003e\u0026thinsp;=\u0026thinsp;12\u0026ndash;20 m. Moreover, in both monoculture and mixed stands, the spatial correlation of the shade-intolerant group relative to the shade-tolerant group showed patterns similar to those of the shade-tolerant group relative to the shade-intolerant group (Fig.\u0026nbsp;\u003cspan refid=\"Fig5\" class=\"InternalRef\"\u003e5\u003c/span\u003ed-\u003cspan refid=\"Fig5\" class=\"InternalRef\"\u003e5\u003c/span\u003ef).\u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003c/div\u003e"},{"header":"4. Discussion","content":"\u003cdiv id=\"Sec15\" class=\"Section2\"\u003e \u003ch2\u003e4.1 Plantations Exhibit Aggregated Distribution after Near-Natural Transformation\u003c/h2\u003e \u003cp\u003eAggregated distribution is the most common spatial pattern in nature (Salas et al., \u003cspan citationid=\"CR101\" class=\"CitationRef\"\u003e2006\u003c/span\u003e; Getzin et al., \u003cspan citationid=\"CR37\" class=\"CitationRef\"\u003e2008\u003c/span\u003e; Wiegand et al., \u003cspan citationid=\"CR128\" class=\"CitationRef\"\u003e2017\u003c/span\u003e), and is also an important trend during plantation succession (Li et al., \u003cspan citationid=\"CR59\" class=\"CitationRef\"\u003e2021c\u003c/span\u003e; Yang et al., \u003cspan citationid=\"CR136\" class=\"CitationRef\"\u003e2025\u003c/span\u003e). Other studies have shown that plantation spatial patterns are influenced not only by ecological processes such as regeneration, self-thinning, and external disturbances, but also jointly regulated by abiotic conditions including topography, soil, and light conditions (Li et al., \u003cspan citationid=\"CR55\" class=\"CitationRef\"\u003e2021a\u003c/span\u003e; Matsushita et al., \u003cspan citationid=\"CR71\" class=\"CitationRef\"\u003e2025\u003c/span\u003e), thereby shifting toward non-regular patterns, which is consistent with our results. In the \u003cem\u003eM. macclurei\u003c/em\u003e and \u003cem\u003eM. laosensis\u003c/em\u003e plantations, planted trees show aggregation at small scales (Appendix 1), and an important reason is the occurrence of sprouting (Yang et al., \u003cspan citationid=\"CR136\" class=\"CitationRef\"\u003e2025\u003c/span\u003e), which supports Hypothesis 1. Sprouting is a strategy by many woody plants to maintain their niches under competition or disturbance, and its occurrence is closely related to species functional traits (Bond and Midgley, \u003cspan citationid=\"CR9\" class=\"CitationRef\"\u003e2001\u003c/span\u003e; Clarke et al., \u003cspan citationid=\"CR17\" class=\"CitationRef\"\u003e2013\u003c/span\u003e). The drivers of sprouting may differ among ecosystems and species (Pausas et al., \u003cspan citationid=\"CR87\" class=\"CitationRef\"\u003e2016\u003c/span\u003e), and our understanding of how sprouting occurs in planted trees and what factors influence it in plantations remains limited (Shi et al., \u003cspan citationid=\"CR109\" class=\"CitationRef\"\u003e2025\u003c/span\u003e). However, it is certain that plantation species that undergo strong differentiation at the root during the first planting are rare. Competition drives the mortality of some trees (Zhang et al., \u003cspan citationid=\"CR146\" class=\"CitationRef\"\u003e2020\u003c/span\u003e), but the extent of pattern change is also influenced by differences in functional traits of planted trees (Forrester et al., \u003cspan citationid=\"CR33\" class=\"CitationRef\"\u003e2018\u003c/span\u003e; Zambrano et al., \u003cspan citationid=\"CR138\" class=\"CitationRef\"\u003e2020\u003c/span\u003e), and these complex relationships further cause the spatial patterns of planted trees to deviate from regularity. In Pingxiang, which is also in the southern subtropical region, the spatial pattern of planted trees in a Chinese fir mixed plantation shifts from a regular pattern at establishment to an aggregated pattern after 27 years of succession (Li et al. \u003cspan citationid=\"CR55\" class=\"CitationRef\"\u003e2021a\u003c/span\u003e, \u003cspan citationid=\"CR59\" class=\"CitationRef\"\u003ec\u003c/span\u003e), supporting our results. Changes in plantation spatial patterns are closely related to planting design, species composition, and management duration.\u003c/p\u003e \u003cp\u003eThe aggregated distribution of regeneration precisely supports Hypothesis 1. Both planted trees have reached physiological maturity and continuously provide seed sources for regeneration, and their capsule and aggregate-fruit traits further cause most seeds to fall near parent trees (Appendix 1). Accordingly, regeneration occurs mainly within the neighborhood of planted trees and shows an aggregated pattern (Appendix 2). Dispersal limitation is also considered an important mechanism underlying aggregated patterns in natural forests (Condit et al., \u003cspan citationid=\"CR21\" class=\"CitationRef\"\u003e2000\u003c/span\u003e; Seidler and Plotkin, \u003cspan citationid=\"CR107\" class=\"CitationRef\"\u003e2006\u003c/span\u003e; Szwagrzyk et al., \u003cspan citationid=\"CR117\" class=\"CitationRef\"\u003e2021\u003c/span\u003e). Because plantations typically have clear management rotations and planted trees are often clear-cut at maturity, regeneration following seed production has been less studied (Yang et al., \u003cspan citationid=\"CR136\" class=\"CitationRef\"\u003e2025\u003c/span\u003e). The soil seed bank is another important source of regeneration (Nathan and Muller-Landau, \u003cspan citationid=\"CR77\" class=\"CitationRef\"\u003e2000\u003c/span\u003e; Singh et al., \u003cspan citationid=\"CR111\" class=\"CitationRef\"\u003e2023\u003c/span\u003e), and it may also increase the intensity and spatial scale of regeneration aggregation. In addition, seedlings are highly sensitive and dependent on microhabitat conditions such as light, moisture, soil, and topography (De Lombaerde et al., \u003cspan citationid=\"CR25\" class=\"CitationRef\"\u003e2020\u003c/span\u003e; Sangsupan et al., \u003cspan citationid=\"CR103\" class=\"CitationRef\"\u003e2021\u003c/span\u003e). Environmental filtering therefore promotes regeneration establishment in locally favorable sites, resulting in aggregation.\u003c/p\u003e \u003cp\u003eBoth shade-tolerant and shade-intolerant regeneration groups show aggregated distributions, but their aggregation intensity differs between monoculture and mixed stands. This pattern is widely considered to be related to understory light conditions (Valladares and Niinemets, \u003cspan citationid=\"CR119\" class=\"CitationRef\"\u003e2008\u003c/span\u003e; Klopčič et al., \u003cspan citationid=\"CR51\" class=\"CitationRef\"\u003e2015\u003c/span\u003e). Row planting creates relatively regular canopy gaps in the stand, with alternating distribution of high and low light in the understory, resulting in a patchy light environment (Zhang et al., \u003cspan citationid=\"CR140\" class=\"CitationRef\"\u003e2019\u003c/span\u003e; Schneider et al., \u003cspan citationid=\"CR105\" class=\"CitationRef\"\u003e2021\u003c/span\u003e). Such uneven light conditions cause shade-tolerant and shade-intolerant regenerations to occur in different light environments. Because \u003cem\u003eM. laosensis\u003c/em\u003e has broad leaves and a dense canopy, shade-intolerant regeneration tends to occur in canopy gaps in its monoculture stand. In contrast, \u003cem\u003eM. macclurei\u003c/em\u003e has smaller leaves and a relatively large crown, and its monoculture stand has better light conditions. As a result, both functional groups can establish under the canopy or in gaps (Yang et al., \u003cspan citationid=\"CR136\" class=\"CitationRef\"\u003e2025\u003c/span\u003e). In the Mixed Stand, differences in crown structure between planted trees create vertical stratification and increase within-stand light availability (Pretzsch, \u003cspan citationid=\"CR91\" class=\"CitationRef\"\u003e2014\u003c/span\u003e; Forrester et al., \u003cspan citationid=\"CR33\" class=\"CitationRef\"\u003e2018\u003c/span\u003e). Consequently, both shade-tolerant and shade-intolerant groups can establish in multiple positions, which reduces their aggregation scale and intensity (Yang et al., \u003cspan citationid=\"CR136\" class=\"CitationRef\"\u003e2025\u003c/span\u003e). Overall, planted trees, planting patterns, and spatial configuration influence the occurrence and distribution patterns of regeneration by altering canopy structure and microhabitat conditions.\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec16\" class=\"Section2\"\u003e \u003ch2\u003e4.2 Spatial Correlations Between Planted Trees and Regenerations in Plantations\u003c/h2\u003e \u003cp\u003eThe degree to which regenerations depend on light determines the direction and strength of their association with planted trees, supporting Hypothesis 2. Shade-tolerant groups can maintain growth under low-light conditions, and their regeneration processes are less dependent on forest gaps (Montgomery and Chazdon, \u003cspan citationid=\"CR75\" class=\"CitationRef\"\u003e2001\u003c/span\u003e; Klopčič et al., \u003cspan citationid=\"CR51\" class=\"CitationRef\"\u003e2015\u003c/span\u003e). Therefore, they can utilize the relatively stable light environment beneath the canopies of planted trees. In contrast, shade-intolerant regenerations tend to occur away from planted trees. For example, in \u003cem\u003ePinus massoniana\u003c/em\u003e plantations, high planting density and canopy closure place seedlings under low-light conditions, which reduces their photosynthetic potential and affects growth (Yan et al., \u003cspan citationid=\"CR134\" class=\"CitationRef\"\u003e2019\u003c/span\u003e; Jin et al., \u003cspan citationid=\"CR49\" class=\"CitationRef\"\u003e2024\u003c/span\u003e; Hao et al., \u003cspan citationid=\"CR41\" class=\"CitationRef\"\u003e2025\u003c/span\u003e). Accordingly, during the near-natural transformation of plantations, the light requirements of seedlings can be leveraged by supplementing shade-tolerant seedlings under the canopy or thinning to improve within-stand light conditions (Cheng et al., \u003cspan citationid=\"CR15\" class=\"CitationRef\"\u003e2017\u003c/span\u003e; Su et al., \u003cspan citationid=\"CR115\" class=\"CitationRef\"\u003e2024\u003c/span\u003e), thereby promoting regeneration and growth within forest gaps and accelerating the transition of plantations from single-layer to multi-layered mixed structures (Liu et al., \u003cspan citationid=\"CR63\" class=\"CitationRef\"\u003e2018\u003c/span\u003e). However, some tree species adopt flexible light adaptation strategies during growth. For instance, \u003cem\u003eMichelia macclurei\u003c/em\u003e exhibits strong shade-tolerance during the seedling stage, enabling survival in forest gaps and under tree canopies, while becoming light-preferring in later developmental stages (Li et al., \u003cspan citationid=\"CR59\" class=\"CitationRef\"\u003e2021c\u003c/span\u003e; Yang et al., \u003cspan citationid=\"CR136\" class=\"CitationRef\"\u003e2025\u003c/span\u003e). Therefore, when both shade-tolerant and shade-intolerant groups are considered at the stand level, the spatial correlation between planted trees and regenerations may be diluted (Fig.\u0026nbsp;\u003cspan refid=\"Fig4\" class=\"InternalRef\"\u003e4\u003c/span\u003ea-\u003cspan refid=\"Fig4\" class=\"InternalRef\"\u003e4\u003c/span\u003ec). This also explains why some studies have found that spatial correlations between planted trees and regenerations are not characterized by a single pattern, with its manifestation varying according to stand structure, observation scale, and functional group differentiation (Ali et al., \u003cspan citationid=\"CR1\" class=\"CitationRef\"\u003e2019\u003c/span\u003e; Li et al., \u003cspan citationid=\"CR59\" class=\"CitationRef\"\u003e2021c\u003c/span\u003e; Zhang et al., \u003cspan citationid=\"CR142\" class=\"CitationRef\"\u003e2022\u003c/span\u003e).\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec17\" class=\"Section2\"\u003e \u003ch2\u003e4.3 Scale-dependent Spatial Correlations between Functional Groups in Plantations\u003c/h2\u003e \u003cp\u003eLight resource partitioning and habitat preferences segregate shade-tolerant and shade-intolerant groups (Montgomery and Chazdon, \u003cspan citationid=\"CR75\" class=\"CitationRef\"\u003e2001\u003c/span\u003e; Valladares and Niinemets, \u003cspan citationid=\"CR119\" class=\"CitationRef\"\u003e2008\u003c/span\u003e; Bartkowicz and Paluch, \u003cspan citationid=\"CR5\" class=\"CitationRef\"\u003e2019\u003c/span\u003e), exhibiting negative correlations at small scales (Fig.\u0026nbsp;\u003cspan refid=\"Fig5\" class=\"InternalRef\"\u003e5\u003c/span\u003ea-\u003cspan refid=\"Fig5\" class=\"InternalRef\"\u003e5\u003c/span\u003ec). However, as the observation scale increases, their spatial correlations are dominated by positive correlations, reflecting the mosaic distribution of light patches in plantations. After the distributions of functional groups span both canopy and gap scales, they tend toward complementary coexistence. This distribution pattern differs from interspecific correlations in natural forests, which often become random as the observation scale increases (Getzin et al., \u003cspan citationid=\"CR35\" class=\"CitationRef\"\u003e2006\u003c/span\u003e; Wiegand et al., \u003cspan citationid=\"CR125\" class=\"CitationRef\"\u003e2007\u003c/span\u003e; Cavard et al., \u003cspan citationid=\"CR13\" class=\"CitationRef\"\u003e2011\u003c/span\u003e). Consistently, in a forest-edge planting experiment in southern Sweden (Wistr\u0026ouml;m and Nielsen, \u003cspan citationid=\"CR130\" class=\"CitationRef\"\u003e2014\u003c/span\u003e), they found that vegetation composed of planted trees and regeneration with different shade tolerance showed spatial differentiation along the forest edge 16 years after establishment. Light-demanding pioneer species occupied high-light environments, whereas shade-tolerant trees and shrubs primarily distributed under relatively low-light canopies, strongly supporting our findings. However, spatial correlations between functional groups may be transient, and planting patterns influence future shifts in functional group spatial correlations (Li et al., \u003cspan citationid=\"CR59\" class=\"CitationRef\"\u003e2021c\u003c/span\u003e). In monoculture stands, microhabitat types are limited and functional-group differentiation is constrained (Pretzsch and Zenner, \u003cspan citationid=\"CR95\" class=\"CitationRef\"\u003e2017\u003c/span\u003e), potentially maintaining spatial associations over extended periods. In mixed stands, interactions among multiple species are alleviated (Forrester, \u003cspan citationid=\"CR31\" class=\"CitationRef\"\u003e2014\u003c/span\u003e; Li et al., \u003cspan citationid=\"CR59\" class=\"CitationRef\"\u003e2021c\u003c/span\u003e), may weaken the associations. Furthermore, spatial correlations between functional groups may be regulated by belowground components. Plant-soil feedbacks (PSFs) promote plant coexistence by regulating relative competitive advantages of species in high-light and low-light environments (Xi et al., \u003cspan citationid=\"CR132\" class=\"CitationRef\"\u003e2023\u003c/span\u003e), thereby reducing correlation strength. This suggests that spatial correlations between functional groups in plantations exhibit spatiotemporal effects and are influenced by multiple ecological processes.\u003c/p\u003e \u003c/div\u003e"},{"header":"5. Conclusion","content":"\u003cp\u003eFollowing the transition of plantations toward near-natural transformation, their spatial patterns have undergone significant alterations. The point patterns of trees in \u003cem\u003eM. laosensis\u003c/em\u003e and \u003cem\u003eM. macclurei\u003c/em\u003e plantations deviate from the initial planting patterns. The distribution patterns of these plantations exhibit aggregated distributions, driven jointly by sprouting, seed dispersal limitation, and within-stand light conditions. This suggests that spatial patterns are controlled by multiple ecological processes, with aggregation represents the ultimate spatial form of forest communities. Tree functional traits determine the direction of spatial correlations between planted trees and regenerations. Therefore, the selection of planted species and planting patterns may be key to maintaining stable regeneration in plantations. Because our study includes only two major local afforestation species, the application of additional species and functional trait combinations in near-natural transformation requires further research. Shade-tolerant groups mainly occur beneath canopies, whereas shade-intolerant groups prefer canopy gaps. Light conditions largely determine patterns of interspecific association, making seedling shade tolerance a priority consideration in the near-natural transformation of plantations. Improving interspecific relationships contributes to optimizing stand structure and enhancing functional stability. The spatial patterns and spatial correlations observed in \u003cem\u003eM. laosensis\u003c/em\u003e and \u003cem\u003eM. macclurei\u003c/em\u003e plantations deepen our understanding of near-natural transformation processes, and they may also be applicable to forest management practice.\u003c/p\u003e"},{"header":"Declarations","content":" \u003cp\u003e \u003cstrong\u003eEthics approval and consent to participate:\u003c/strong\u003e \u003cp\u003eNot applicable\u003c/p\u003e \u003cp\u003e \u003cstrong\u003eConsent for publication:\u003c/strong\u003e \u003cp\u003eNot applicable\u003c/p\u003e \u003ch2\u003eCompeting interests:\u003c/h2\u003e \u003cp\u003eAuthors have no conflicts of interest\u003c/p\u003e \u003ch2\u003eFunding:\u003c/h2\u003e \u003cp\u003eThis work was supported the National Natural Science Foundation of China (Grant No. 32060340).\u003c/p\u003e\u003ch2\u003eAuthor Contribution\u003c/h2\u003e\u003cp\u003eYuanfa Li conceived the idea, Ziyi Chen and Yuanfa Li wrote the manuscript, all authors collected dataset from wild and reviewed the manuscript.\u003c/p\u003e\u003ch2\u003eAcknowledgement\u003c/h2\u003e\u003cp\u003eMaster Studies from College of Forestry, Guangxi University investigated dataset in wild, they were Jintao Li, Yongyi Wei, Liting Wei, Xiuqiong Luo.\u003c/p\u003e\u003ch2\u003eData Availability\u003c/h2\u003e\u003cp\u003eThe datasets used and/or analysed during the current study are available from the corresponding author on reasonable request.\u003c/p\u003e"},{"header":"References","content":"\u003col\u003e\n \u003cli\u003eAli A, Dai D, Akhtar K, Teng M, Yan Z, Urbina-Cardona N, Mullerova J, Zhou Z (2019) Response of understory vegetation, tree regeneration, and soil quality to manipulated stand density in a \u003cem\u003ePinus massoniana\u003c/em\u003e plantation. 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Forest Ecology and Management 465: 118103.https://doi.org/10.1016/j.foreco.2020.118103\u003c/li\u003e\n\u003c/ol\u003e"}],"fulltextSource":"","fullText":"","funders":[],"hasAdminPriorityOnWorkflow":false,"hasManuscriptDocX":true,"hasOptedInToPreprint":true,"hasPassedJournalQc":"","hasAnyPriority":false,"hideJournal":false,"highlight":"","institution":"","isAcceptedByJournal":false,"isAuthorSuppliedPdf":false,"isDeskRejected":"","isHiddenFromSearch":false,"isInQc":false,"isInWorkflow":false,"isPdf":false,"isPdfUpToDate":true,"isWithdrawnOrRetracted":false,"journal":{"display":true,"email":"[email protected]","identity":"new-forests","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":false,"externalIdentity":"nefo","sideBox":"Learn more about [New Forests](http://link.springer.com/journal/11056)","snPcode":"11056","submissionUrl":"https://submission.nature.com/new-submission/11056/3","title":"New Forests","twitterHandle":"","acdcEnabled":true,"dfaEnabled":true,"editorialSystem":"em","reportingPortfolio":"Springer Hybrid","inReviewEnabled":true,"inReviewRevisionsEnabled":false},"keywords":"Plantation, Planting pattern, Distribution pattern, Spatial correlation, Functional groups","lastPublishedDoi":"10.21203/rs.3.rs-8701430/v1","lastPublishedDoiUrl":"https://doi.org/10.21203/rs.3.rs-8701430/v1","license":{"name":"CC BY 4.0","url":"https://creativecommons.org/licenses/by/4.0/"},"manuscriptAbstract":"\u003cp\u003eThe spatial pattern of plantations changes after near-natural transformation, but the underlying driving mechanisms remain unclear. To address this issue, we established three permanent plots in monoculture and mixed plantations of native tree species \u003cem\u003eMytilaria laosensis\u003c/em\u003e Lecomte and \u003cem\u003eMichelia macclurei\u003c/em\u003e Dandy in southern China. Trees were classified into planted trees and regenerations, and regenerations were further divided into shade-tolerant and shade-intolerant functional groups based on light adaptability. We applied the univariate pair correlation function \u003cem\u003eg\u003c/em\u003e\u003csub\u003e11\u003c/sub\u003e(\u003cem\u003er\u003c/em\u003e) to analyze the distribution patterns of planted trees and regenerations, and bivariate pair correlation function \u003cem\u003eg\u003c/em\u003e\u003csub\u003e12\u003c/sub\u003e(\u003cem\u003er\u003c/em\u003e) to analyze spatial correlations between planted trees and regenerations, as well as between functional groups. Results indicate: (1) planted trees exhibited aggregated distributions in all stands at small spatial scales (\u003cem\u003er\u003c/em\u003e\u0026thinsp;=\u0026thinsp;0\u0026ndash;2 m, \u003cem\u003er\u003c/em\u003e\u0026thinsp;=\u0026thinsp;0\u0026ndash;3 m), regenerations showed aggregation across a wider range of spatial scales; (2) spatial correlations between planted trees and regenerations were positive in the \u003cem\u003eMytilaria laosensis\u003c/em\u003e Lecomte monoculture stand and the mixed stand (\u003cem\u003er\u003c/em\u003e\u0026thinsp;=\u0026thinsp;0\u0026ndash;2 m), while it was negatively correlated in monoculture stand of \u003cem\u003eMichelia macclurei\u003c/em\u003e Dandy (\u003cem\u003er\u003c/em\u003e\u0026thinsp;=\u0026thinsp;2\u0026ndash;20 m); (3) spatial correlations between shade-tolerant and shade-intolerant regenerations was negatively correlated in all stands, but at slightly different scales (\u003cem\u003er\u003c/em\u003e\u0026thinsp;=\u0026thinsp;0\u0026ndash;2 m, \u003cem\u003er\u003c/em\u003e\u0026thinsp;=\u0026thinsp;0\u0026ndash;3 m, \u003cem\u003er\u003c/em\u003e\u0026thinsp;=\u0026thinsp;0\u0026ndash;5 m). These results suggest that planting patterns and functional traits jointly drive spatial pattern shifts in plantations, leading to increased spatial complexity within stands and deepening our understanding of the near-natural transformation process of plantations.\u003c/p\u003e","manuscriptTitle":"Spatial Patterns of Michelia macclurei and Mytilaria laosensis plantations after 24 Years of Near-Natural Transformation","msid":"","msnumber":"","nonDraftVersions":[{"code":1,"date":"2026-04-10 19:52:50","doi":"10.21203/rs.3.rs-8701430/v1","editorialEvents":[{"type":"communityComments","content":0},{"type":"reviewersInvited","content":"","date":"2026-04-06T17:13:15+00:00","index":"","fulltext":""},{"type":"editorAssigned","content":"","date":"2026-01-28T07:36:42+00:00","index":"","fulltext":""},{"type":"checksComplete","content":"","date":"2026-01-28T07:29:37+00:00","index":"","fulltext":""},{"type":"submitted","content":"New Forests","date":"2026-01-26T14:25:36+00:00","index":"","fulltext":""}],"status":"published","journal":{"display":true,"email":"[email protected]","identity":"new-forests","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":false,"externalIdentity":"nefo","sideBox":"Learn more about [New Forests](http://link.springer.com/journal/11056)","snPcode":"11056","submissionUrl":"https://submission.nature.com/new-submission/11056/3","title":"New Forests","twitterHandle":"","acdcEnabled":true,"dfaEnabled":true,"editorialSystem":"em","reportingPortfolio":"Springer Hybrid","inReviewEnabled":true,"inReviewRevisionsEnabled":false}}],"origin":"","ownerIdentity":"8ae14e11-b915-49ab-8154-6b21fbdbea64","owner":[],"postedDate":"April 10th, 2026","published":true,"recentEditorialEvents":[],"rejectedJournal":[],"revision":"","amendment":"","status":"under-review","subjectAreas":[],"tags":[],"updatedAt":"2026-04-10T19:52:50+00:00","versionOfRecord":[],"versionCreatedAt":"2026-04-10 19:52:50","video":"","vorDoi":"","vorDoiUrl":"","workflowStages":[]},"version":"v1","identity":"rs-8701430","journalConfig":"researchsquare"},"__N_SSP":true},"page":"/article/[identity]/[[...version]]","query":{"redirect":"/article/rs-8701430","identity":"rs-8701430","version":["v1"]},"buildId":"XKTyCvWXoU3ODBz1xrDgd","isFallback":false,"isExperimentalCompile":false,"dynamicIds":[84888],"gssp":true,"scriptLoader":[]}