1. Introduction
Plant communities are dynamic, characterized by seasonal fluctuations, interannual changes, and succession [1,2]. Community dynamics is an important aspect of ecological studies [3], providing crucial information on changes in community structure and composition over time. Quantitative or attribute changes in species composition, abundance, diversity, and community structure are considered indicators of community dynamics [4,5,6,7].
The research method of community dynamics is an important topic in community ecology, and various methods are able to assess different aspects of community dynamics [8]. For example, community characteristics are monitored by field observations [9], and trends in community change can be predicted via model simulations [10]. Stages of community development are often evaluated by comparing the characteristics and relationships of communities with different origins [11]. Community establishment processes are usually determined by analyzing the dominance state of a target species in different communities [12].
Among the various methods for characterizing community dynamics, comparing the composition and status of dominant species determines the community trajectory [12]. Understanding the relationship between the community and local environment is helpful for explaining facets of community performance [13,14], while analyzing stability is critical for quantifying community dynamics and the factors influencing it [15,16]. Integrating these methods enables a better description of community characteristics and stability of a given target species.
Forest ecosystems play an important role in biodiversity conservation, ecosystem services, and climate regulation [2]. Ecological research on forests with different tree species is of great significance. For example, many Picea species have attracted much research attention, such as P. asperata [17,18], P. purpurea [19,20], and P. crassifolia [21,22]. Picea jezoensis (PJ) is an important evergreen coniferous tree species in Northeast Asia [23,24]. Many ecological studies have focused on PJ, including its regeneration characteristics [25,26,27,28], growth–environment relationships [29,30,31,32], community and diversity [33,34], and the impacts of climate change [35]. These studies have enriched our understanding of PJ’s ecology in places such as Korea, Japan, and Far East Russia. In particular, the stand structure of PJ forests is key research theme at the community level [33,34].
Besides the natural distribution area of PJ mentioned above, China is also an important distribution area of PJ; this species has been the subject of many ecological studies. Most of these focused on radial growth and ecological factors [36,37,38,39,40,41], community and diversity [42,43,44,45,46], regeneration and growth characteristics [47,48,49,50], species–environment relationships [51,52], and the impacts of climate change [53]. That body of research has arguably increased our understanding of the ecological characteristics of PJ in China. Community ecology studies have mainly addressed the species composition and community structure of PJ forests in China [42,43,44,45].
Yet, to our best knowledge, few studies have investigated the community dynamics of PJ [33,45,46,47], leaving several fundamental questions unaddressed, thereby limiting our knowledge of the establishment and changes in the PJ community and hindering the effective protection and restoration of PJ forests. For example, what is the threshold for changes in the dominance position of PJ in the community? Since the PJ community can exhibit regional differences [34], a detailed analysis of the characteristics and dynamics of the PJ community in a specific area is needed. Studying this question provides a scientific basis for the conservation of local PJ forests, as well as the formulation of targeted sustainable forest management strategies.
To answer the above scientific question, this study attempted to propose the hypothesis based on the knowledge accumulation and research experience in forest ecology. Dominant species in forest communities are those with a large number of individuals, extensive crown cover, and high biomass [2,3]. This study presumed that a shift in the dominance status of PJ from being the second dominant species to the first dominant species in a community would occur incrementally. This study hypothesized that this shift would happen when the dominance degree of PJ reached a critical threshold value.
A survey of forests with PJ trees in three counties of Northeast China was conducted to investigate the PJ community’s characteristics and dynamics. The objectives were to (1) describe the dominant species composition and environment of the PJ community and (2) determine the threshold for a dominance shift of PJ in the community. By comparing the different forests containing PJ trees, the aim was to comprehensively assess the PJ community and gauge changes in PJ’s dominance status to provide empirical information for the conservation of PJ forests, especially in protecting existing PJ forests and restoring degraded PJ forests.
2. Materials and Methods
2.1. Study Area and Data Collection
This field survey was conducted in Antu, Fusong, and Changbai counties of Jilin Province, China. Antu County, in eastern Jilin, encompasses 7444 km2, and has 86% forest cover, with a wood volume of 98 million m3, where the annual average temperature is 2.2 °C in the south and 3.6 °C in the north, whose corresponding annual average precipitation is 669.7 and 594.7 mm [54]. Fusong County (6159 km2) is located in southeastern Jilin, and has 86.7% forest cover, with a wood volume of 4.62 million m3, where the annual average temperature is 4 °C and the annual average precipitation is 800 mm [55]. Changbai County is also in southeastern Jilin and has an area of 2506 km2. Its forest cover is 92%, with a wood volume of 23.34 million m3; the annual average temperature is 3.9 °C, and the annual average precipitation is 605.8 mm [56].
The forests in the study area consist of various tree species. For example, the top seven ranked tree species in terms of relative basal area (RBA) are the following: PJ (RBA = 0.264), Abies nephrolepis (AN) (RBA = 0.143), Larix olgensis (LO) (RBA = 0.101), Pinus koraiensis (PK) (RBA = 0.088), Populus ussuriensis (PU) (RBA = 0.086), Betula costata (BC) (RBA = 0.082), and Tilia amurensis (TA) (RBA = 0.080).
The forest survey was carried out in 2014. Forty-eight forest plots (each 600 m2) containing PJ trees (23 plots in Antu County, 22 plots in Fusong County, and 3 plots in Changbai County) were surveyed (Figure 1; Table S1). The distance between any two plots was ≥4 km. In every plot, the scientific name and stem size of all living trees with a diameter at breast height (DBH) ≥ 5 cm were recorded. A total of 2635 trees were measured. The basic information of each plot was also recorded, namely, its elevation, slope aspect, slope degree, and forest origin (Table S1). The elevation of the 48 plots ranged from 840 to 1810 m, and all plots were located at sites with a gently slope (≤10°). All plots were located in natural forests.
2.2. Data Analysis
Species composition is the most important factor determining the attributes of a community [2]; comparing the species composition of different communities with PJ trees highlights the characteristics of PJ communities. The environment has an impact on the composition and structure of communities [3]; analyzing the relationship between different communities with PJ trees and environmental factors reveals the environmental conditions of PJ community. On the basis of clarifying the PJ community characteristics and its environmental conditions, quantifying the dominance state and change of PJ in the community helps to understand the PJ community dynamics in a simple and convenient way.
2.2.1. Species Composition of Different Forests Containing P. jezoensis Trees
This study analyzed the dominant species composition of different forests with PJ trees. The RBA of each tree species per plot was calculated, and the dominant species identified accordingly [12,16]. RBA was calculated by dividing the basal area of a species by the total basal area of all species [57]. Dominant species were defined as those with an RBA > 0.10 [12,16]. In each plot, the first two dominant species and their RBA values were listed, and PJ’s rank was determined. The plots were divided into four groups based on the PJ’s ranking and dominance: (i) PJ as the first dominant species, (ii) PJ as the second dominant species, (iii) PJ as the third (or less ranked) dominant species, and (iv) PJ as a non-dominant species. Next, the dominant species composition and proportions in the groups were analyzed and compared. To do that, the species name and proportion of the first two dominant species in each group were determined, and then the dominant species that co-appeared among the groups were identified. This study focused on PJ and the top two dominant species and sought to highlight the dominant species composition of the PJ community (i.e., PJ as the first dominant species).
2.2.2. Environmental Characteristics of Different Forests Containing P. jezoensis Trees
This study examined the local environment of different forests with PJ trees. Slope is a complex factor affecting the tree species and their growth status [2]. Although there were differences in slope aspect and degree among the sample plots, all were nonetheless situated at gently sloping sites (Table S1). So, we considered that 48 plots had the same slope conditions.
Tree species are usually only distributed within a certain elevational range, and vegetation type and species composition may vary with elevation [2]. Because the elevation span of the 48 plots was about 1000 m, the relationship between forest and elevation cannot be ignored. Hence, the elevational characteristics of different forests with PJ trees were described and compared to highlight the role of elevation for the PJ community.
To determine the distribution pattern of forests with PJ trees vis-à-vis elevation, we calculated the mean elevation of each group and tested whether elevation differed significantly among the four groups using one-way analysis of variance (ANOVA) or a non-parametric test based on the statistical requirements [58]. If so, we further detected differences between any two groups using multiple comparisons; either the least significant difference (LSD) or Games–Howell method was used, based on the homogeneity of variance [59].
2.2.3. Changes to the Dominance Status of P. jezoensis
Temporary stability (TS) was employed as a robust metric to assess the dominance dynamics of PJ, providing insights into the critical threshold where shifts in dominance occur. The TS provides a measure of forest community stability [15,16] and was calculated this way:
where d1 and d2 denote the dominance of the first and second ranked species, respectively [15,16]. This study used basal area for dominance. The top two ranked species in dominance have the same basal area when TS = 0, which corresponds to the threshold for a change in the first dominant species.As seen above, TS conveys the relationship between the top two ranked species in dominance. Therefore, only plots in groups where PJ was the first or second dominant species were retained for this analysis. To determine the feasibility of combining the plots from two groups for a unified analysis, the environmental characteristics of these two groups were evaluated. All plots were located at gently sloping sites (Table S1), so these two groups thus had the same conditions in slope. Then, the t test or non-parametric test [58] was used to determine whether elevation differed significantly between the two groups. Furthermore, the dominant species composition of these two groups was also described and compared.
This study analyzed the relationship between the dominance status and degree of dominance of PJ in the community to determine this critical value. TS and the RBA were used to represent the dominant state and the degree of dominance of PJ, respectively. These indicators were calculated for the retained plots. According to its definition [15], if TS has a large value, this indicates that the first dominant species is less likely to be replaced by PJ in plots where it is the second dominant species, and PJ is less likely to be replaced by the second dominant species in plots where PJ is the first dominant species. To integrate these two groups of plots into a unified research system, TS values for plots where PJ was the second dominant species were converted to negative values. The correlation between TS and the RBA of PJ was analyzed to determine whether or not they exhibited a linear trend. If so, to verify whether the linear relationship meets the statistical requirements, ANOVA, the determination coefficient (R2), significance level, and standardized residuals were used to assess the regression model [60]. The threshold RBA of PJ was calculated by setting TS = 0.
3. Results
3.1. Dominant Species Composition of the P. jezoensis Community
Based on the ranking and dominance characteristics of PJ, the 48 plots were divided into four groups. BC and AN mainly constituted the second dominant species in plots where PJ was ranked the first dominant species (13 plots) (Figure 2; Table S2). BC, AN, and PK mainly constituted the first dominant species for plots in which PJ was the second dominant species (14 plots) (Figure 2; Table S3). PU and LO mainly constituted the first dominant species for plots where PJ was a co-dominant but not ranked among the top two dominant species (8 plots) (Figure 2; Table S4). In plots where PJ was not the dominant species (13 plots), LO was the first dominant species in 30.77% of the plots, while PU, QM, and TA had the same proportions (15.38%) (Figure 2; Table S5).
Among the four groups, three species (PK, PU, and LO) co-appeared as the first dominant species in the three groups in which PJ was not the first dominant species. Two species (PK and PU) co-appeared as the second dominant species in the group in which PJ was the first dominant species (Figure 2; Tables S2–S5).
3.2. Distribution of P. jezoensis Community Across Elevation
Different forests with PJ trees exhibited a regular distribution with respect to elevation. Forests in which PJ was the first dominant species were distributed at the highest elevation, with a mean value of 1408 (±198) m (Figure 3).
Elevation differed significantly among the four groups (ANOVA, F = 5.549, p < 0.01). Based on the multiple comparisons testing, there were no significant differences between adjacent groups (such as groups 1 vs. 2, groups 2 vs. 3, and groups 3 vs. 4) (p > 0.05), but there were significant differences between separated groups (such as groups 1 vs. 3, groups 1 vs. 4, and groups 2 vs. 4) (p < 0.05; Figure 3; Table 1).
To sum up, the PJ community occupied sites at high elevation (mean: 1408 m), this being similar to the mean elevation of the community in which PJ was the second dominant species, yet significantly higher than those of communities in which PJ was co-dominant but not among the top two dominant species or not a dominant species.
3.3. Dominance States and Changes of P. jezoensis in the Forest Community
Plots in which PJ was not one of the first two dominant species were excluded from the subsequent analysis. A total of 27 plots were retained for this analysis, comprising 13 plots in which PJ was the first dominant species and 14 plots in which PJ was the second dominant species.
In terms of their environmental conditions, both the 13 plots and 14 plots were located at sites with a gentle slope degree of ≤10°. The mean elevation of the 13 plots was 1408 (±198) m, which exceeded 1311 (±241) m of the 14 plots, albeit not significantly (t test, t = 1.134, df = 25, p > 0.05).
Regarding species composition, the second dominant species of the 13 plots consisted of BC, AN, PU, FM, and PK, of which four (BC, AN, PU, and PK) co-appeared as the first dominant species in the 14 plots (Figure 2; Tables S2 and S3).
The TS was strongly correlated with the RBA of PJ (r = 0.944; p < 0.001). In the fitted linear model, the R2 was 0.890, the p-value of the ANOVA was < 0.001 (F = 202.884), and the standardized residuals ranged from –2.074 to 2.028. Thus, a strong, predictable linear relationship existed between the TS and the RBA of PJ. The regression model’s equation was TS = 2.383 × RBA − 0.922. Based on this model, when TS = 0, the RBA of PJ = 0.387 (Table 2; Figure 4).
4. Discussion
4.1. P. jezoensis Community and Environmental Characteristics
Comparing the dominance of a target species in different communities is capable of revealing the species’ community characteristics [12]. Based on the PJ ranking and dominance characteristics, the 48 surveyed plots were divided into four groups. The results show that PJ exhibits different dominance states in different forests, a feature similar to that of a fir species of Abies faxoniana [12]. The employed method of community grouping analysis was able to reveal the dominance characteristics of target species in different forests.
Different species have different roles in a given community [3]. From a species composition perspective, evidently BC and AN were the main tree species in the PJ community, while PK and PU were the co-appeared species of PJ. This result is mostly consistent with findings of previous studies [42,53]. Some studies have analyzed the relationship of PJ to other tree species [61,62]. However, the data of the present study did not lend support to such species association and correlation analyses; hence, further studies are needed to reveal the actual relationship between these tree species and PJ. Furthermore, the dominant species differed among forests with PJ trees, indicating that PJ is capable of establishing and growing in a variety of forest types and that suitable PJ habitats are diverse. Alongside PJ’s wide distribution range in terms of elevation (Table S1), our results support a previous study that reported PJ has a wide niche breadth [63].
In mountainous conditions, vegetation distribution is usually strongly related to elevation [2]. Although PJ’s distribution does range widely across elevation, PJ was not always the first dominant species in every forest community [44]. For example, communities in which PJ is the first dominant species, this being the typical PJ community, prevailed at the highest elevation with a mean value of 1408 m. The first dominant species (PJ) in the PJ community was unlike those of the other three community groups that occupied different elevations. Given that the forest origins of all surveyed plots were natural forest processes, the mean elevation of 1408 m is a pivotal parameter for describing the natural distribution of the PJ community. In this way, the distributional elevation of the PJ community in this study is more specific than previous descriptions for this species [42,44]. Furthermore, many previous studies have analyzed the response of PJ’s growth to climate change [37,39,41,49], additional studies should be considered to reveal the potential impact of climate change on the distributional elevation of PJ.
4.2. P. jezoensis Community Stability and State Change
Many studies on community stability have been conducted, focusing on the definition of stability [64,65], diversity–stability relationships [66,67], and influential factors [68,69]. Other studies have assessed the PJ community using a subordinate function and spatial structure index [70,71]. However, no empirical studies have yet explored the PJ community’s stability. This study, the first to do so, revealed a linear relationship between TS and the RBA of PJ. Importantly, based on this relationship, the critical value when PJ changes from being the second dominant species to the first dominant species was an RBA = 0.387. This key result fills a research gap in quantifying the shift in the dominance status of PJ in the forest community and provides useful information for studying the community dynamics of other forest types.
Under largely consistent slope conditions and similar elevation, this study found that PJ exhibited a different ranking and dominance status and that four species co-appeared as the dominants in forests where PJ was the first and second dominant species. We conjecture that the PJ ranking and dominance state are reversible. Considering the PJ community occupied the highest elevation of 1408 m, this could be the most suitable elevation for the growth of PJ trees. Therefore, the higher the elevation, the more likely a site is to be conducive to improving PJ’s ranking and dominance status. This phenomenon requires further observation and verification, however.
Many indicators are used to gauge forest community stability [69,70,71,72,73]. As a recently proposed indicator, TS has been adopted to analyze the community stability of forests in Northeast and Southwest China [15,16], but its scope of application should still be determined. In this study, our results confirm that TS can be used to analyze and detect the shift in dominance status from the second to the first dominant species. However, this indicator could be improved to accommodate a wider range of community dynamics.
4.3. Limitations and Implications
This study described the characteristics and dynamics of the PJ community. Nonetheless, it has some limitations. First, this work is based on a single survey; long-term continuous observations should therefore be considered. Second, the sample size was 48 plots; using a larger sample size would provide even more comprehensive information. Third, this study described the characteristics of the PJ community from the perspective of trees; incorporating the understory vegetation would facilitate a more detailed description. Fourth, this study depicted the PJ community mainly in relation to elevation and slope. We suggest that future studies incorporate other plausible influencing factors, such as soil [70], wind [74], and insects [75]. Fifth, due to differences in environment and tree species, the linear model and critical value obtained in this study may not necessarily have broad applicability, so it should be used cautiously in other areas and with other tree species.
By assessing the dominant species composition and environmental characteristics of the PJ community, this study offers an approach to studying the community ecology of a targeted plant species. We analyzed the relationship between the dominance status and the degree of dominance to obtain the critical value when PJ changed from being the second dominant species to the first dominant species, thus providing a quantitative method to objectively analyze changes in a plant community. PJ is not only distributed in China but also occurs in regions of Northeast Asia. Furthermore, many other tree species are growing in forests with PJ trees. Accordingly, the research ideas and methods from this study can provide a valuable reference for studying the PJ forests outside of China, as well as the community ecology of other tree species.
The forests of Northeast China play a fundamental role in timber reserves, biodiversity conservation, mitigating climate change, and ecological functions [76], so protecting these forests is of great significance. This study provides a scientific basis for the conservation of PJ forests. For instance, in this study, PK and PU co-appeared as the first dominant species in the three groups where PJ was not the first dominant species, and they also served as the second dominant species in the group where PJ was the first dominant species. That is, the status of PJ in communities dominated by these two species changed from no advantage to a slight advantage to a strong advantage and to replacing these species. The establishment of the PJ community consisted of replacing the first dominant species with PJ, providing potential paths for restoring PJ forests.
Furthermore, the derived elevation distribution provides location information for the protection or restoration site selection of PJ forest tracts. The biotic (such as species composition) and abiotic (such as slope and elevation) factors reported on here can provide environmental information for restoring PJ forest. The critical value threshold is helpful for judging the attributes and status of forest communities. Moreover, this study’s findings also provide comparable information for ecological research and conservation of PJ forests in other areas in its natural range across Northeast Asia.
5. Conclusions
Based on its ranking and dominance characteristics, Picea jezoensis (PJ) exhibited differing states of dominance in different forests surveyed. By comparing the forests containing PJ trees, we find that the PJ community occupies gentle sites at high elevations, and that tree species co-occurring with PJ mainly included Betula costata and Abies nephrolepis. The linear relationship between temporary stability (TS) and the relative basal area (RBA) of PJ explained its dominance status shift in the local community. To facilitate greater comprehensive understanding, studying the dynamics of more PJ populations in broader areas during long-term surveys, while also considering more soil, geographical, and biotic factors, is strongly suggested.
Conceptualization, D.K.; methodology, D.K.; formal analysis, J.D., Z.J. and D.K.; investigation, S.G. and Y.L.; writing—original draft preparation, J.D.; writing—review and editing, D.K.; supervision, J.L.; funding acquisition, J.D. and D.K. All authors have read and agreed to the published version of the manuscript.
Not applicable.
The original contributions presented in the study are included in the article/
The authors declare no conflicts of interest.
Footnotes
Disclaimer/Publisher’s Note: The statements, opinions and data contained in all publications are solely those of the individual author(s) and contributor(s) and not of MDPI and/or the editor(s). MDPI and/or the editor(s) disclaim responsibility for any injury to people or property resulting from any ideas, methods, instructions or products referred to in the content.
Enlarge this image.Figure 2. Composition of the first or second dominant species of different forests containing Picea jezoensis (PJ) trees (group 1: PJ was the first dominant species; group 2: PJ was the second dominant species; group 3: PJ was a co-dominant but not ranked among the top two dominant species; group 4: PJ was not the dominant species; 1a, 2a, 4f: Betula costata (BC); 1b, 2b, 4e: Abies nephrolepis (AN); 1c, 2d, 3a, 4b: Populus ussuriensis (PU); 1d: Fraxinus mandshurica (FM); 1e, 2c, 3d, 4g: Pinus koraiensis (PK); 2e, 3b, 4a: Larix olgensis (LO); 3c: Betula platyphylla (BP); 3e, 4d: Tilia amurensis (TA); 4c: Quercus mongolica (QM)).
Enlarge this image.Figure 3. Mean elevation of different forests containing Picea jezoensis (PJ) trees (group 1: PJ was the first dominant species; group 2: PJ was the second dominant species; group 3: PJ was a co-dominant but not ranked among the top two dominant species; group 4: PJ was not the dominant species; a, b, c: bars that shared a letter (i.e., a, b, or c) do not differ significantly from one another).
Enlarge this image.Figure 4. Temporary stability (TS) and relative basal area (RBA) of Picea jezoensis (PJ).
Multiple comparisons (using LSD method) of elevation among different forests containing Picea jezoensis (PJ) trees (group 1: PJ was the first dominant species; group 2: PJ was the second dominant species; group 3: PJ was a co-dominant but not ranked among the top two dominant species; group 4: PJ was not the dominant species).
| Item | Mean Elevation Difference (m) | Standard Error | p Value |
|---|---|---|---|
| Group 1–Group 2 | 97 | 81 | 0.239 |
| Group 1–Group 3 | 234 | 95 | 0.017 |
| Group 1–Group 4 | 314 | 83 | 0.000 |
| Group 2–Group 3 | 138 | 93 | 0.148 |
| Group 2–Group 4 | 217 | 81 | 0.010 |
| Group 3–Group 4 | 80 | 95 | 0.405 |
Linear regression model between temporary stability (TS) and the relative basal area (RBA) of Picea jezoensis (PJ).
| Model | Unstandardized Coefficient (UC) | Standard Error | t Value | p Value | 95% Confidence Interval of UC |
|---|---|---|---|---|---|
| RBA | 2.383 | 0.167 | 14.244 | 0.000 | [2.039, 2.728] |
| Constant | −0.922 | 0.078 | −11.793 | 0.000 | [−1.083, −0.761] |
Supplementary Materials
The following supporting information can be downloaded at:
References
1. Shang, Y. General Ecology; 3rd ed. Peking University Press: Beijing, China, 2010; pp. 338-362.
2. Li, J.; Niu, S.; Liu, Y. Forest Ecology; 3rd ed. Higher Education Press: Beijing, China, 2017; pp. 17–32, 95–118, 208–254, 312–351.
3. Niu, C.; Lou, A.; Sun, R.; Li, Q. Foundations in Ecology; 4th ed. Higher Education Press: Beijing, China, 2023; pp. 72–100, 160–201.
4. He, J.N.; Ning, C.W.; Zhang, W.T.; ÜmÜt, H.; Shen, Z.H. Altitudinal patterns of population structure dynamics of Picea schrenkiana in the northern slope of the eastern Tianshan Mountains. Acta Ecol. Sin.; 2024; 44, pp. 5389-5398.
5. Li, J.Y.; Chen, X.X.; Li, M.W.; Zhang, N.N.; Zhou, Z.Q.; Li, X.M.; Ding, J.L.; He, Q.H.; Shi, F.S. Community composition and structure under different restoration pathways in subalpine forest, Sichuan Province, China. Acta Ecol. Sin.; 2024; 44, pp. 3471-3482.
6. Niu, Y.D.; Cai, T.J. Changes in species diversity and influencing factors in secondary forest succession in northern Da Hinggan Mountains. Chin. J. Plant Ecol.; 2024; 48, pp. 349-363.
7. Tian, Z.; Wang, Y.; Duan, W.; Wang, Z.; Yang, W.; Chen, L. Study on spatial structure characteristics of broadleaved Pinus koraiensis forest at different succession stages in the Xiaoxing’an Mountains. J. Southwest For. Univ. (Nat. Sci.); 2024; 44, pp. 130-137.
8. Sun, Z.; Zhou, D. Ecology Research Methods; Science Press: Beijing, China, 2010; pp. 120-131.
9. Li, J.; Li, X.Y.; Cao, J.; Tan, L.Z.; Zhao, X.H. Community structure and dynamics of secondary coniferous and broad-leaved mixed forest in Changbai Mountains. Acta Ecol. Sin.; 2020; 40, pp. 1195-1206.
10. Zhang, W.W.; Yin, D.H.; Lei, Y.T.; Han, X.T.; Peng, J.Y.; Zhao, Z. Structural dynamics and succession simulation of the dominant populations in three types of Quercus acutissima community in the southern Loess Plateau. Acta Ecol. Sin.; 2024; 44, pp. 2572-2581.
11. Ouyang, Z.Y.; Li, Z.X.; Liu, J.G.; An, L.; Zhang, H.M.; Tan, Y.C.; Zhou, S.Q. The recovery processes of giant panda habitat in Wolong Nature Reserve, Sichuan, China. Acta Ecol. Sin.; 2002; 22, pp. 1840-1849.
12. Kang, D.; Lv, J.; Li, S.; Chen, X.; Wang, X.; Li, J. Community change characteristics of Abies faxoniana: A case study in the Wanglang Nature Reserve. Ecol. Indic.; 2019; 107, 105594. [DOI: https://dx.doi.org/10.1016/j.ecolind.2019.105594]
13. Lu, J.Y.; Zhang, C.Y.; Zhao, X.H. Relationship between forest strata structure and topography of confierous and broad-leaved mixed forest in Jiaohe City, Jilin Province. Acta Ecol. Sin.; 2021; 41, pp. 6613-6620.
14. Tong, Y.W.; Qu, L.L.; Fu, Q.X.; Chen, Y.B.; Xiang, X.Y.; Zhu, W.D.; Qi, G.; Dai, L.M. Species diversity of forest plant communities on the southern slope of the Dabie Mountains and its relationship with altitude factors. Acta Ecol. Sin.; 2024; 44, pp. 5307-5317.
15. Jia, Z.; Ge, S.; Li, Y.; Kang, D.; Li, J. Relationship between tree richness and temporary stability of plant communities: A case study of a forest in Northeast China. Forests; 2021; 12, 1756. [DOI: https://dx.doi.org/10.3390/f12121756]
16. Kang, D.; Li, J. Temporary stability characteristics and influencing factors of Abies faxoniana-dominated communities in the Wanglang Nature Reserve. Forests; 2022; 13, 1186. [DOI: https://dx.doi.org/10.3390/f13081186]
17. Liu, Q.; Yin, H.J.; Wu, Y. A study on the structure of Picea asperata comunity of Miyaluo subalpine in Western Sichuan, China. J. Mt. Sci.; 2003; 21, pp. 695-701.
18. Yang, X.; Li, J.; Fan, N.; Wang, Y.; Zhang, Z. Effects of gap size on natural regeneration in Picea asperata forests of Northern China. Forests; 2023; 14, 2102. [DOI: https://dx.doi.org/10.3390/f14102102]
19. Zhao, Z.; Kang, D.; Guo, W.; Zhao, L.; Cui, L.; Li, J. Climate sensitivity of purple cone spruce (Picea purpurea) across an altitudinal gradient on the eastern Tibetan Plateau. Dendrochronologia; 2019; 56, 125586. [DOI: https://dx.doi.org/10.1016/j.dendro.2019.03.006]
20. Zhao, Y.; Cao, J.; Li, B.; Qi, R.; Liu, T.; Chen, X.; Gao, B.; Zhao, Y.; Liu, J.; Cao, X. Niche of woody plant populations of Picea purpurea community in Dayugou forest area, Taohe Nature Reserve, Gansu Province. Acta Ecol. Sin.; 2022; 42, pp. 1865-1875.
21. He, Z.; Zhao, W.; Liu, H.; Zhang, Z. Successional process of Picea crassifolia forest after logging disturbance in semiarid mountains: A case study in the Qilian Mountains, northwestern China. For. Ecol. Manag.; 2010; 260, pp. 396-402. [DOI: https://dx.doi.org/10.1016/j.foreco.2010.04.034]
22. Zhao, W.J.; Liu, X.D.; Jing, W.M.; Xu, L.H.; Niu, Y.; Qi, P.; Zhao, Y.H. Spatial heterogeneity of community structure of Picea crassifolia forest in Qilian Mountains, China. Chin. J. Appl. Ecol.; 2015; 26, pp. 2591-2599.
23. Aizawa, M.; Yoshimaru, H.; Saito, H.; Katsuki, T.; Kawahara, T.; Kitamura, K.; Shi, F.; Sabirov, R.; Kaji, M. Range-wide genetic structure in a north-east Asian spruce (Picea jezoensis) determined using nuclear microsatellite markers. J. Biogeogr.; 2009; 36, pp. 996-1007. [DOI: https://dx.doi.org/10.1111/j.1365-2699.2008.02074.x]
24. Thomas, P.; Zhang, D.; Katsuki, T.; Rushforth, K. Picea jezoensis. The IUCN Red List of Threatened Species 2013: e.T42325A2972665. Available online: https://www.iucnredlist.org/species/42325/2972665 (accessed on 16 May 2024).
25. Nakagawa, M.; Kurahashi, A.; Hogetsu, T. The regeneration characteristics of Picea jezoensis and Abies sachalinensis on cut stumps in the sub-boreal forests of Hokkaido Tokyo University Forest. For. Ecol. Manag.; 2003; 180, pp. 353-359. [DOI: https://dx.doi.org/10.1016/S0378-1127(02)00654-0]
26. Iijima, H.; Shibuya, M. Evaluation of suitable conditions for natural regeneration of Picea jezoensis on fallen logs. J. For. Res.; 2010; 15, pp. 46-54. [DOI: https://dx.doi.org/10.1007/s10310-009-0133-9]
27. Yano, K.; Shibuya, M. Site preference and occurrence patterns of Picea jezoensis and Abies sachalinensis on decayed logs in natural coniferous forests in Hokkaido, northern Japan. J. For. Res.; 2010; 15, pp. 108-114. [DOI: https://dx.doi.org/10.1007/s10310-009-0162-4]
28. Han, A.R.; Kim, H.J.; Jung, J.B.; Park, P.S. Seed germination and initial seedling survival of the subalpine tree species, Picea jezoensis, on different forest floor substrates under elevated temperature. For. Ecol. Manag.; 2018; 429, pp. 579-588. [DOI: https://dx.doi.org/10.1016/j.foreco.2018.07.042]
29. Han, A.R.; Lee, S.K.; Suh, G.U.; Park, Y.; Park, P.S. Wind and topography influence the crown growth of Picea jezoensis in a subalpine forest on Mt. Deogyu, Korea. Agric. For. Meteorol.; 2012; 166–167, pp. 207-214. [DOI: https://dx.doi.org/10.1016/j.agrformet.2012.07.017]
30. Jang, W.; Keyes, C.R.; Running, S.W.; Lim, J.H.; Park, P.S. Climate growth relationships of relict Picea jezoensis at Mt. Gyebang, South Korea. For. Sci. Technol.; 2015; 11, pp. 19-26.
31. Ukhvatkina, O.; Omelko, A.; Zhmerenetsky, A. Local topography has significant impact on dendroclimatic response of Picea jezoensis and determines variation of factors limiting its radial growth in the Southern Sikhote-Alin. Forests; 2023; 14, 2050. [DOI: https://dx.doi.org/10.3390/f14102050]
32. Ukhvatkina, O.N.; Omelko, A.M.; Zhmerenetsky, A.A. Changes in the dendroclimatic response of the Picea jezoensis (Siebold & Zucc.) Carriere along altitudinal gradient in the southern Sikhote-Alin. Contemp. Probl. Ecol.; 2023; 16, pp. 745-757.
33. Jang, W.; Park, P.S. Stand structure and maintenance of Picea jezoensis in a northern temperate forest, South Korea. J. Plant Biol.; 2010; 53, pp. 180-189. [DOI: https://dx.doi.org/10.1007/s12374-010-9103-1]
34. Park, B.J.; Byeon, J.G.; Heo, T.I.; Cheon, K.; Yang, J.C.; Oh, S.H. Comparison of species composition among Picea jezoensis (Siebold & Zucc.) carrière forests in Northeast Asia (from China to South Korea). J. Asia-Pac. Biodivers.; 2023; 16, 272e281.
35. Park, C.; Byeon, J.; Park, J.; Lee, S.; Park, B.; Kim, J.; Woo, H. Impact of climate change on species distribution of Picea jezoensis Carrière in Baekdudaegan using ensemble modeling. Sens. Mater.; 2022; 34, pp. 4629-4637. [DOI: https://dx.doi.org/10.18494/SAM4163]
36. Yu, D.; Wang, Q.; Wang, G.G.; Dai, L. Dendroclimatic response of Picea jezoensis along an altitudinal gradient in Changbai Mountains. Sci. China Ser. E Technol. Sci.; 2006; 49, pp. 150-159. [DOI: https://dx.doi.org/10.1007/s11434-006-8116-0]
37. Li, G.Q.; Bai, F.; Sang, W.G. Different responses of radial growth to climate warming in Pinus koraiensis and Picea jezoensis var. komarovii at their upper elevational limits in Changbai Mountain, China. Chin. J. Plant Ecol.; 2011; 35, pp. 500-511.
38. Wang, S.L.; Zhao, Y.; Gai, X.R.; Wang, X.Y.; Yu, D.P.; Zhou, W.M.; Zhou, L.; Dai, L.M. Response of radial growth of Picea jezoensis var. komarovii to climate factors along an altitudinal gradient on Changbai Mountain, Northeast China. Chin. J. Ecol.; 2017; 36, pp. 3131-3137.
39. Yu, J.; Liu, Q.J.; Zhou, G.; Meng, S.W.; Zhou, H.; Xu, Z.Z.; Shi, J.N.; Du, W.X. Response of radial growth of Pinus koraiensis and Picea jezoensis to climate change in Xiaoxing’anling Mountains, Northeast China. Chin. J. Appl. Ecol.; 2017; 28, pp. 3451-3460.
40. Zhu, L.; Cooper, D.J.; Yang, J.; Zhang, X.; Wang, X. Rapid warming induces the contrasting growth of Yezo spruce (Picea jezoensis var. microsperma) at two elevation gradient sites of northeast China. Dendrochronologia; 2018; 50, pp. 52-63.
41. Gong, X.; Xie, R.; Yang, H. Response of radial growth of three common tree species to climate change in a spruce-fir mixed stand in Changbai Mountain of northeastern China. J. Beijing For. Univ.; 2023; 45, pp. 1-10.
42. Chen, L. Preliminary study on the structure of Picea jezoensis forest on the southwest slope of Changbai Mountain. Ser. Plant Ecol. Geobot.; 1963; 1, pp. 69-80.
43. Shao, B.; Deng, K.M. The composition and important value of plant species in subalpine spruce-fir forest on the northern slope of Changbai Mts. J. Nat. Resour.; 2000; 15, pp. 66-73.
44. Jiang, P.; Zhao, G.; Ye, J.; Cui, G.; Deng, H. Structure of forest communities on the northern slope of Changbai Mountain and its variation along elevation gradients. Chin. J. Ecol.; 2003; 22, pp. 28-32.
45. Lei, X.; Lu, Y.; Peng, C.; Zhang, X.; Chang, J.; Hong, L. Growth and structure development of semi-natural larch-spruce-fir (Larix olgensis–Picea jezoensis–Abies nephrolepis) forests in northeast China: 12-year results after thinning. For. Ecol. Manag.; 2007; 240, pp. 165-177. [DOI: https://dx.doi.org/10.1016/j.foreco.2006.12.019]
46. Chen, K.Y.; Zhang, H.R.; Zhang, B.; He, Y.J. Population structure and dynamic characteristics of typical constructive species in natural secondary forest on the northern slope of Changbai Mountain. Acta Ecol. Sin.; 2021; 41, pp. 5142-5152.
47. Deng, K.M.; Shao, B.; Li, F. Analysis for diameter height structures and growth of Spruce-fir forest on northern slope of Changbai Mountains. Resour. Sci.; 1999; 21, pp. 77-84.
48. Yu, Z.L.; Yu, G.R.; Wang, Q.F.; Zhao, S.D.; Yan, X.D. Canopy gap characteristics and their influences on the regeneration of broad-leaved Pinus Koriensis forests in Changbai Mountain. Resour. Sci.; 2001; 23, pp. 64-68.
49. Qin, X. Growth response of main tree species to climate change and disturbance in an ecotone on Changbai Mountains. J. Northeast. For. Univ.; 2015; 43, pp. 42-45.
50. He, D.N.; Yang, H.; Wen, J.; Xie, R. Density and spatial distribution of seedlings and saplings in different gap sizes of a spruce-fir mixed stand in Changbai Mountains, China. Chin. J. Appl. Ecol.; 2020; 31, pp. 1916-1922.
51. Bu, R.C.; Chang, Y.; Hu, Y.M.; Li, X.Z.; He, H.S. Sensitivity of coniferous trees to environmental factors at different scales in the Small Xing’an Mountains, China. J. Plant Ecol.; 2008; 32, pp. 80-87.
52. Liu, Q.; Bi, L.; Song, G.; Wang, Q.; Jin, G. Species-habitat associations in an old-growth temperate forest in northeastern China. BMC Ecol.; 2018; 18, 20. [DOI: https://dx.doi.org/10.1186/s12898-018-0177-9] [PubMed: https://www.ncbi.nlm.nih.gov/pubmed/29986704]
53. Zhang, P.Y.; Liang, Y.; Ma, T.X.; Liu, B.; Wu, M.M. The migration of main tree species lags behind climate change in Changbai Mountains. Chin. J. Ecol.; 2022; 41, pp. 1674-1682.
54. Antu County Government. 2022; Available online: http://www.antu.gov.cn/xq/dldm/202204/t20220414_377913.html (accessed on 24 May 2024).
55. Fusong County Government. 2024; Available online: http://www.fusong.gov.cn/lyfs/fsgk/zrdl/201706/t20170612_96794.html (accessed on 24 May 2024).
56. Changbai County Government. 2023; Available online: http://www.changbai.gov.cn/cbgk/zrzy/201801/t20180108_257152.html (accessed on 24 May 2024).
57. Fang, J.; Wang, X.; Shen, Z.; Tang, Z.; He, J.; Yu, D.; Jiang, Y.; Wang, Z.; Zheng, C.; Zhu, J. et al. Methods and protocols for plant community inventory. Biodivers. Sci.; 2009; 17, pp. 533-548.
58. Yan, H.; Xu, Y.; Zhao, N.; Yang, S.; Wang, T. Medical Statistics; 3rd ed. People’s Medical Publishing House: Beijing, China, 2015; pp. 444-457.
59. Xie, L.H.; Shang, T. SPSS Statistical Analysis and Data Mining; Publishing House of Electronics Industry: Beijing, China, 2012; pp. 163-188.
60. Backhaus, K.; Erichson, B.; Plinke, W.; Wang, X.; Weiber, R. Multivariate Statistical Analysis: Using SPSS Tools; Truth and Wisdom Press: Shanghai, China, Shanghai People’s Publishing House: Shanghai, China, 2009; pp. 35-80.
61. Ni, R.Q.; Tang, J.Y.; Cheng, Y.X.; Zhang, C.Y.; Chen, B.B.; He, H.J.; Xia, F.C. Spatial distribution patterns and associations of main tree species in spruce-fir forest in Changbai Mountains, northeastern China. J. Beijing For. Univ.; 2013; 35, pp. 28-35.
62. Qiang, Y.Q.; Fan, C.Y.; Zhang, C.Y. Species diversity maintenance mechanism of dark coniferous forests community in Changbai Mountain. Acta Ecol. Sin.; 2023; 43, pp. 1884-1891.
63. Liu, W.; Cao, W. Niche characteristics of main plant species in spruce-fir forests in Changbai Mountains. Chin. J. Ecol.; 2011; 30, pp. 1766-1774.
64. Pimm, S.L. The complexity and stability of ecosystems. Nature; 1984; 307, pp. 321-326. [DOI: https://dx.doi.org/10.1038/307321a0]
65. Ives, A.R. Diversity and stability in ecological communities. Theoretical Ecology: Principles and Applications; 3rd ed. May, R.; Mclean, A.; Tao, Y.; Wang, B. Higher Education Press: Beijing, China, 2010; pp. 116-129.
66. McCann, K.S. The diversity–stability debate. Nature; 2000; 405, pp. 228-233. [DOI: https://dx.doi.org/10.1038/35012234] [PubMed: https://www.ncbi.nlm.nih.gov/pubmed/10821283]
67. Zhang, J.H.; Huang, Y.M. Biodiversity and stability mechanisms: Understanding and future research. Acta Ecol. Sin.; 2016; 36, pp. 3859-3870.
68. Guo, Q.Q.; Zhang, W.H.; Cao, X.P. Establishment of an evaluation model of the forest community stability based on fuzzy synthetic evaluation: A case study of main forest communities in Huanglong Mountains. Sci. Silvae Sin.; 2009; 45, pp. 19-24.
69. Song, W.; Zhang, H. Research status and hot spot analysis of forest ecosystem stability based on CNKI. For. Sci. Technol.; 2022; 3, pp. 48-52.
70. Zhang, M.T.; Zhang, Q.; Kang, X.G.; Yang, Y.J.; Xu, G.; Zhang, L.X. Community stability for spruce-fir forest at different succession stages in Changbai Mountains, Northeast China. Chin. J. Appl. Ecol.; 2015; 26, pp. 1609-1616.
71. Zhou, M.L.; Zhang, Q.; Kang, X.G.; Guo, W.W.; Ji, L.; Li, C.F. Study on the stability of forest communities based on the spatial structure index. Plant Sci. J.; 2016; 34, pp. 724-733.
72. Zheng, Y.R. Comparison of methods for studying stability of forest community. Sci. Silvae Sin.; 2000; 36, pp. 28-32.
73. Zhang, M.X.; Wang, D.X.; Peng, S.L.; Huang, Y.K.; Zhang, G.G. Community stability analysis for the oak-pine mixed forest in Qinling Mountains. Acta Ecol. Sin.; 2015; 35, pp. 2564-2573.
74. Wang, H.Y.; Zhang, Y.J.; Jin, Y.H.; Xu, J.W.; Tao, Y.; He, H.S.; Gao, X.; Bai, Y.Y.; Hu, R.; Han, Y.Y. Vegetation damage, post-disaster change degree and driving factors in the cold temperate wind disaster area of Changbai Mountain, China. Chin. J. Appl. Ecol.; 2019; 30, pp. 1580-1588.
75. Yu, D.J.; Zheng, Y.G.; Sun, C.H.; Zhao, C.; Jin, Y.H.; Zhang, L.J.; Li, J.G.; Liu, L.J. Investigation and factors analysis of Dendrolimus superans outbreaks in Changbai Mountain National Nature Reserve. For. Res.; 2022; 35, pp. 103-111.
76. National Forestry and Grassland Administration. China Forest Resources Report; China Forestry Publishing House: Beijing, China, 2019; pp. 78-241.
© 2024 by the authors. Licensee MDPI, Basel, Switzerland. This article is an open access article distributed under the terms and conditions of the Creative Commons Attribution (CC BY) license (https://creativecommons.org/licenses/by/4.0/). Notwithstanding the ProQuest Terms and Conditions, you may use this content in accordance with the terms of the License.