Soil physicochemical properties during vegetation restoration
Our results showed that soil BD decreased, and the contents of SOC, TN, TP, TK, Ca, AN, and AP increased with vegetation restoration (Fig. 2), indicating that soil physicochemical properties improved significantly. These results are partially consistent with our hypothesis and with the results of Zhang et al. (2019).
The rapid recovery of SOC at our study site has been proven to be affected by plant biomass and soil nutrients (Gu et al. 2019). The response rate of SOC (140.76%) in this research was higher than the results under semi-arid conditions (71%) recorded by Boix-Fayos et al. (2009), which may be due to the more humid conditions in subtropical regions. Consistent with the rapid accumulation of SOC, the rates of change in TN and AN were greater than the SOC change. This result differs from the results of studies in the same subtropical area of southwest China (Xu et al. 2018), which may be due to differences in the degree of degradation and type of vegetation system. Additionally, soil N is also input from other N sources, such as atmospheric N deposition, and symbiotic N fixation by legumes (Alday et al. 2012). This explains why the recovery rates of TN and AN were greater than SOC. Our results for the increase in TP and AP contents are consistent with the results of Zhang et al. (2019), who proposed that soil TP and AP contents gradually increase with the composition of tree species, annual litter yield, and SOC content along with the development of a forest’s second succession. This is also supported by the significant positive correlation of soil TP and AP contents with the species diversity, biomass, and contents of SOC, TN and AN observed in this study (Fig. 5), which suggests that the accumulation of SOC improves soil nutrients during vegetation restoration (Zhang et al. 2019).
The variation ranges of BD and pH in the subtropical regions of China are 0.97–1.47 g·cm− 3 and 4.5–6.0, respectively (Hunan Provincial Department of Agriculture 1989). Our results were in the variation ranges. All the pH samples in our study indicate that soil pH (4.54–4.96) was lower than the results of Takoutsing et al. (2016), being formed by a moderate ferrallitic effect under high temperature and high humidity conditions in subtropical regions (Li et al. 2012b). Meanwhile, decreasing soil BD and pH has also been attributed to the accumulation of organic matter, which is conducive to the formation of soil aggregates and the improvement of soil microbial activity (Bienes et al. 2016). This in turn releases a large number of small molecular organic acids during the decomposition of organic matter (van Breemen et al. 1984), resulting in a decline in soil BD and pH values. The SOC in our study increased and showed negative relationships with BD and pH during vegetation restoration (Fig. 5). In addition, this study shows that biomass stimulated the decrease in soil pH during vegetation restoration (Fig. 3). The accumulation of biomass led to increased biomass in the roots, almost certainly reflecting the development of the vegetation community from annual plant species to perennial plants, which is more conducive to the release and accumulation of the various acid exudates (Pang et al. 2018). Although BD and pH showed a general declining trend, their values reached a peak in the 10–12 years restoration period (Fig. 2). These results may be caused by a combination of factors (i.e. soil texture, vegetation types and soil acid–base equilibrium). Firstly, as herbs developed into shrubs in our study site, the erosion effect of rainwater on soil silt and clay particles resulted in a high proportion of sand particles in the 10–12 years restoration period, reflecting the transformation of soil texture to sandy soil with high BD (Wang et al. 2018b). Secondly, changes in vegetation types could be a major driver behind the difference in cations absorption of the vegetation and consequent variation in the proportions of soil cations (Gu et al. 2019). From soil acid–base equilibrium, the increase in cations (especially Mg and AK) suggests that the soil H+ was replaced by increased alkaline ions (Berthrong et al. 2009). Due to the similar soil parent materials at different restoration stages, soil Ca, Mg, and K contents, which are all derived from parent rock materials, change little in response rates during vegetation restoration (Takoutsing et al. 2016).
Vegetation development during restoration periods
In our study, species diversity increased with an 86.36% recovery rate as restoration progressed, and these results are consistent with the results of Wang et al. (2018a). The amount of biomass increased significantly with the greatest recovery rate (2906.52%) over the old growth forest, followed by H (596%) and DBH (128%) in the 45–46 years period. These results are partially consistent with our hypothesis and are very similar to the results of Hu et al. (2017).
Improvements to the soil environment can provide community habitat quality which then promotes the enrichment of community diversity (Huang et al. 2015). Ca content had a significant positive effect on species diversity (Fig. 3), reflected in the following mechanisms. Firstly, Ca2+ has the function of maintaining the homeostasis of intracellular ions, especially in acidic soil where higher Ca2+ content can counterbalance the toxicity of aluminum ions for plants, further improving plant resistance to adversity and being conducive to the improvement of community diversity (Roem et al. 2002). Secondly, the increase in soil Ca content alongside vegetation restoration can be instrumental in the coexistence of species with different Ca requirements and the settlement of calcium-loving species (Hooper 1998). Additionally, soil P determines the species composition of a vegetation community (Huang et al. 2015); thus, soil AP content was considered as another major factor determining species diversity increase (Fig. 3).
In our study, biomass, DBH, and H had basically the same changing trend, and were all significantly affected by SOC, TN, AN, and AP contents (Figs. 4 and 5). This is consistent with Brandies et al. (2009), who demonstrated that there are significant positive growth rates and similar effect factors between biomass, DBH, and H in a general case. Data analysis of our study site confirmed that the percentages of individual trees with DBH greater than 8 cm and H greater than 5 m were larger in the 45–46 years restoration period (54% and 77% respectively) than in the old growth forest (41% and 63% respectively) (Chen et al. 2019). The greatest values of DBH and H in the 45–46 years restoration period may be because Pinus massoniana, as the dominant species, is a fast-growing heliophilous plant that gets more light by increasing vertical growth (H) (Cheng et al. 2011).
Soil SOC, TN, AN, and AP contents were leading factors in stimulating the increase in biomass, DBH, and H (Fig. 3). As the environmental basis for vegetation survival, improving the soil provides a better habitat and essential nutrients for vegetation growth (Huang et al. 2018), ultimately promoting the positive succession of vegetation (Liang et al. 2010). The accumulation of SOC affects biomass, DBH, and H mainly by decomposing and releasing large amounts of nutrients to meet plant growth needs, and by improving soil texture and promoting microbial activity which provide a better growing environment for vegetation (Alday et al. 2012). Moreover, the increase in N content promotes growth of the leaf area and improves plant photosynthesis, providing sufficient energy for the growth of individual plants. P is the nutrient that most limits productivity and species richness (Huang et al. 2015), and also controls leaf litter decomposition (Zeng et al. 2016). In addition, soil P changes the structure of the root system, promotes the formation and growth of fine roots, lateral roots and secretions of root exudates, and thereby stimulates plants to make more efficient use of soil nutrients (Li et al. 2017).
Key factors affecting vegetation restoration
Soil factors (pH, SOC, TN, TK, Mg, AN, and AP) and vegetation features (biomass, DBH, and H) were the main factors influencing vegetation restoration at our study site. This is consistent with the finding that the recovery of degraded ecosystems not only relies on soil rehabilitation, but also on the reconstruction, productivity, and function of vegetation (Liang et al. 2010; Peng et al. 2012).
The soil properties and vegetation features can be viewed as three distinct groups. The first group can be summarized as soil pH, SOC, TN, AN, AP, biomass, DBH, and H across the vegetation restoration periods. The roles of soil pH, SOC, TN, AN, and AP have been analyzed above. Specifically, soil resource is the main limiting factor in the early period of vegetation restoration. However, in the later period of vegetation restoration, the change in community characteristics leads to light conditions becoming a limiting factor (van Der Maarel and Franklin 2013). With the accumulation of biomass, a complex community structure reduces the understory light transmittance, controlling the vegetation in the understory including the growth and mortality of tree seedlings and saplings (Montgomery and Chazdon 2001). Therefore, the shade tolerant species are successively established, increasing understory vegetation richness. On the other hand, heliophilous species are shaped by increasing H and diameter to gain more light by adapting to strong interspecific competition. At increasingly larger H and DBH, light transmittance could further influence a species’ light-capturing ability and distribution (Cheng et al. 2011). The limitation of light conditions for vegetation growth and performance in the late vegetation restoration period means that increases in biomass, DBH, and H are the key growth factors, determining restoration success.
The second group of soil variables includes Mg. The increase in Mg during the restoration periods was accompanied by a series of improvements in the plants’ physiological processes, such as photosynthetic efficiency, carbohydrate metabolism, and synergistic absorption with P (Unger 2010). The third group showed that the vegetation restoration development was determined by TK. Besides N and P, K is the limiting nutrient with a significant influence on vegetation growth and development (Pang et al. 2018), mainly reflected in the impact on plant photosynthesis and respiration by controlling the regulation of stomata opening (Unger 2010), even though here the effect of TK on vegetation development was not significant.
Previous studies have suggested that species diversity was the dominant vegetation factor for vegetation restoration in a large scale (Crouzeilles et al. 2016), because higher species richness can enhance ecosystem stability and increase nutrient use efficiency (Hu et al. 2017). However, species diversity was not considered to be an influential factor for vegetation restoration in our study. The difference could be due to the non-significance of the relationships between species diversity and the main soil physicochemical properties or biomass, indicating that species diversity had no significant effect on the recovery of soil fertility and plant communities at our study site. In addition, species diversity showed a decreasing trend in the 45–46 years restoration period (Fig. 2), in which dominant species transformed from simple shrubs and herbs to pioneer species such as Pinus massoniana. In fact, needles of some Pinus species have been reported as a hindering factor which influences the regeneration of native plants and increases in species diversity (Navarro-Cano et al. 2010). It is reasonable that species diversity has no significant effect on vegetation restoration in specific study area, but further research is needed.
Soil and vegetation factors affecting biomass
The variation of biomass was one of the important indexes reflecting vegetation restoration (Mansourian et al. 2005). Therefore, the relative importance of soil properties and vegetation features in driving biomass development can reflect the degree of their individual and joint influence on vegetation restoration.
Our study revealed that the change in biomass was strongly influenced by the interaction of soil properties and vegetation features, which explained 55.55%–72.32% of the biomass variation (Fig. 4). This dominant contribution by joint influence to biomass may be explained by the close interaction between vegetation and soil (Liang et al. 2010). As we discussed above, there was a clear co-evolutionary relationship between soil factors (pH, SOC, TN, AN, and TP) and vegetation features (DBH and H) across the restoration periods. This result suggests that the variations in key soil factors (pH, SOC, TN, AN, and TP) were likely to promote the growth of plant and the restoration of vegetation structure (Alday et al. 2012). In turn, vegetation features (DBH, and H) could influence improvements in the soil environment (Fig. 3). These results also offer the further evidence for the hypothesis that the mechanisms of plant and soil promote vegetation restoration synergistically.
This study also found that soil properties explained 3.30%–31.44% of the variation in biomass, which was basically higher than explanation of vegetation (5.09%–24.32%). This result provides evidence that the importance of soil properties in driving the changes observed in biomass is more than that of vegetation features in the study region, which is most likely because the advantage of hydrothermal conditions in the subtropical region accelerates the material circulation, and promotes the enrichment of soil organic matter (Corlett and Hughes 2015); thus, providing a fertile environment for plant growth. The regulation mechanism of soil properties on biomass development had been discussed previously. With vegetation restoration, the increased in plant species has intensified the competition of aboveground parts for light resources and underground roots for soil resources (Cheng et al. 2011; Li et al. 2017), which further induces the variations of individual growth and morphological structure of trees (DBH and H). As DBH and H increased, more fine materials and litters can be intercepted and accumulated by plants, further enhancing the accumulation of biomass (Li et al. 2017).
The biomass development at our study site was influenced by different soil and vegetation factors in different restoration periods. In the early restoration period (4–5 years), SOC was the major influential factor (Table 4). The possibility is that SOC is the main source of most nutrients, and that the accumulation of SOC promotes improvements in other soil factors, such as TN, AN, and AP, which have a notable effect on vegetation growth and development (Alday et al. 2012). In the 4–5 years restoration period, SOC content was low (Fig. 2), which is not conducive to the improvement of soil structure or the accumulation of nutrients (Bienes et al. 2016). Therefore, the low SOC not only limits the growth of plant roots, but also intensifies the contradiction between the demand of plant growth for water and nutrients and the supply of soil water and nutrients, resulting in hindrances to plant growth.
H, pH, and AP were the main factors driving biomass development in the 10–12 years restoration period. This could be attributed to the competition of shrubs for light, which would drive the increasing of H to adapt to interspecific competition (Cheng et al. 2011). Additionally, the accumulation of biomass impels plants to need more N- and P-rich substances (such as enzymes, transport proteins, and amino acids) to participate in metabolic activities (Qin et al. 2016). Therefore, shrubs need to absorb more N and P for growth than do herbs. In particular, P is an important limited factor in red soil area of south China (Gao et al. 2014). However, in the 10–12 years restoration period, the increasing of pH affected the availability of P (Duan et al. 2008), suggesting that the role of AP may intensity the inequity of competition among plants, rather than promote the accumulation of biomass.
Biomass in the 45–46 years restoration period was conditioned by the synergistic effect of H and pH. The significant effects of H and pH may be caused by a combination of two factors. Firstly, the dominant tree species (Pinus massoniana) of 45–46 years restoration period obtains more light by increasing H and canopy density (Cheng et al. 2011), resulting in lower density of woody plants (Table 1); thus, H had a negative effect on biomass. Secondly, low soil pH is beneficial to improve soil permeability, aggregates and porosity (such as BD), and the accumulation of soil nutrients (such as SOC, N and P) (Ramírez et al. 2015), and enhances the availability of P, K, Ca, and Mg (Duan et al. 2008). Meanwhile, soil pH decreased with vegetation restoration, and the bioaccumulation and material circulation increased with advantageous hydrothermal conditions (Corlett and Hughes 2015), which were beneficial to the increment of soil nutrient content; thus restoration stimulates the increase of biomass.
In the old growth forest (sub-climax community), the structure of plant community has reached a state of stable (Peng et al. 2012), which means that the development of vegetation features (DBH and H) has entered a slow growth stage and has less of an impact on biomass. Instead, as a nutrient bank and soil health indicator (Bienes et al. 2016), SOC continues to influence biomass growth. In addition, evergreen trees with a long leaf life need to accumulate more organic substances (such as lignin) to construct defensive structures, and require higher N and P content to maintain normal growth and metabolism (Zeng et al. 2016). Therefore, the supply capacity of soil N and P largely determines the effectiveness of vegetation restoration (Li et al. 2012a).
