1. Introduction
Moso bamboo (
Phyllostachys pubescens (Pradelle) Mazel ex J.Houz.) is extensively distributed in south-eastern and southern Asia, important to the global carbon cycle and water conservation, and also an important forest species because it grows fast and has a short life cycle. Bamboo is well known for rapid accumulation of biomass, high annual output of timber, and as a source of food in the form of tender bamboo shoots. Improving the operating efficiency and economic returns from bamboo forests is, therefore, important, which among other things, requires suitable methods to examine and monitor bamboos growing in forests—methods that are efficient, feasible, and accurate. Traditional ways of managing forests of pure bamboo have resulted in the decline of the sites of such forests, lower soil fertility, and increasing damage by pests and diseases, whereas many recent studies have shown that mixed forests comprising bamboo and broad-leaved trees are conducive to maintaining ecosystem balance, improving stand structure, and increasing soil fertility [
1]. Studying the relationship between broad-leaved trees and bamboo in such mixed forests will help to manage them efficiently and to obtain higher economic returns.
The growth of individual trees, whether in pure stands or mixed stands, is affected by many factors and by the interactions between those factors, including climate, site conditions, the ontogenetic stage, and competition [
2]. The structure of plant communities is shaped by competitive interactions among individual plants [
3,
4]. In a forest, these interactions determine the size and the position of individual canopies in the overall forest canopy, which affects light interception, photosynthesis, growth, and the survival of individual plants [
5]. Thus, competition among canopies of individual trees significantly affects virtually every benefit we derive from forest ecosystems [
6].
In establishing new forests, it is easy to choose a pattern in which a few broad-leaved trees are surrounded by bamboo. The canopy of broad-leaved trees is the dominant layer in mixed forests of bamboo and broad-leaved trees, and competition between these two components means that one bamboo or broad-leaved tree affects the viability and growth of another. The shape of the crown and the photosynthetic efficiency of a broad-leaved tree and the space it occupies can change the competitive ability of a bamboo plant. Crown characteristics provide insights into the process of absorption and consumption of nutrients by individual trees, which also affect their competitive ability. An index of competition based on canopy can reflect such competition between trees better than the indexes based on other characteristics [
7]. The spatial structures of bamboos and broad-leaved trees in mixed forests are more distinct than those in other types of forest.
Field measurements provide the largest and most important dataset for forest surveys, and the assessment of tree metrics is essential in evaluating forests. However, such measurements require a great deal of time and labour, which is why automated methods of collecting such data are urgently required. One such method, namely terrestrial laser scanning (TLS)—which is efficient, objective, non-destructive, and accurate—has been used extensively. The method also makes it possible to acquire three-dimensional (3D) data, inventories, and data on canopy characteristics [
8,
9,
10,
11,
12,
13]. Many studies show that point clouds obtained from TLS can extract such details as the diameter at breast height (DBH) and at other heights, the positions of individual trees [
14,
15], and trunk volume [
16]. In addition, trunk biomass [
12], leaf area index [
17], and the surface area and volume of the canopy can also be estimated using TLS [
18], which gives results accurate to the nearest millimetre [
19]. Compared with the cumbersome traditional forestry surveys, which cannot accurately extract the information of the tree canopy, the application of TLS provides a new option for measuring the size and shape of crown [
20]. Compared to the cumbersome traditional forestry surveys, which cannot offer any reliable estimates of the tree canopy, TLS offers an easier way for measuring the size and the shape of a tree crown quickly and accurately.
It is against this background that the present study sought to quantify the crowns of broad-leaved trees in mixed forests and to assess the effect of the canopy on the growth of bamboo [
11,
13]. More specifically, the study aimed at (1) constructing an index of competition between broad-leaved trees and bamboos based on the canopy characteristics of the broad-leaved trees and (2) analysing this competition relationship between broad-leaved trees and bamboos.
4. Discussion
Competition is one of the most basic relationships between individuals [
6], and intraspecific competition is normally greater than interspecific competition [
24], which suggests that competition between bamboos should be greater than that between bamboos and broad-leaved trees—results from the present study provide an insight into how the latter influenced the growth of bamboo. We found that broad-leaved trees had suppressed the growth of bamboo growing closer than 3 m, but promoted it when the distance was greater than 3 m up to the point at which it exceeded the range of influence of the broad-leaved trees (
Figure 3 and
Figure 4,
Table 5). As we observed, bamboos with the largest DBH were found 3 m away from broad-leaved trees. Due to the cover provided by the canopy of broad-leaved trees, bamboo cannot grow normally with insufficient lighting, when the distance is less than 3 m because it cannot get enough sunlight. The average DBH of bamboo increases initially and then stabilizes at distances farther than 3 m. The amount of photosynthetically active radiation can characterize the intensity of competition well [
25]. There is no parameterization for canopy characteristics of broad-leaved trees and bamboo. Therefore, the current study provides new tools to evaluate these effects. Because the broad-leaved trees formed canopies that permitted more sunlight to filter through them, it compensated to some extent for the light blocked by the overlapping canopies and thus promoted the growth of bamboo [
26]. These observations suggest that the main factor affecting the growth of broad-leaved trees and bamboo is light. Under the canopy of broad-leaved trees, bamboo showed almost no growth, except under gaps in the canopy. However, when sufficient light was available, the litter of broad-leaved trees serves as a source of nutrients for bamboo growing around the edges of the canopy; as a result, bamboo grows better. Therefore, in analysing the competition between bamboo and broad-leaved trees, we need to take into account not only the spatial structure of the canopy but also the availability of nutrients below the soil surface.
Differences in crown shapes, canopy efficiency, and the extent of space occupied change the intensity of competition among trees [
27]. The survival environment and the living space occupied by an individual tree are good indicators of competition. The three most important factors that determine the extent of competitive pressure exerted by the canopy on the nearby trees are (1) canopy size, which determines the size of the overlapping area and the projected area, (2) tree height, which determines whether the canopy can project into the space to cast a shadow over nearby plants, and (3) the distance between adjacent canopies, which directly determines the degree of competition between trees.
The competition index based on features of the canopy correlated better with the crown than the simple model did [
26,
28]. In many tree species, the DBH is closely correlated to tree height and crown shape [
29]. Compared to the DBH, features of the canopy (tree height, projected area, etc.) have less influence on the competition whereas the DBH is often directly related to the competitive ability of a tree. The diameter and distance, which are used by the distance-dependent Hegyi competition index, are easy to measure, and the results are highly reliable. However, the overlap volume between the canopies of broad-leaved trees and of bamboo is also a matter of distance in some sense. Therefore, a competition index that considers the canopy is also a distance-dependent competition index. In the present study, the greater overlapping volume proved better at explaining the degree of competition between broad-leaved tree and bamboo.
Because the sampling spots were selected at random and the sample was small, we could not assess the joint impact of the entire forest. The competition between canopies of a single broad-leaved tree and bamboo has not been studied; therefore, we cannot evaluate the competition between different tree species and bamboo. These two sources of uncertainty have hindered the analysis of the competition between bamboo and broad-leaved trees in mixed forests. Although our data can quantify the effects of broad-leaved trees on the growth of bamboo, their competing relationships cannot be explained at the species level. The effects of different species of broad-leaved trees on bamboo may have superimposed effects. The effects of different species of broad-leaved trees on bamboo may be more complex. Therefore, future research on the competition between bamboo and broad-leaved trees in mixed forests should take into account different tree species to devise better models or at least devise models that are specific to a given tree species and bamboo.
As we found, the complex forest environment led some errors in extraction of TLS point cloud data, which make the average recognition rate of individual bamboo lower than 1, as shown in
Figure 5.
A reasonable number and locations of scanning stations will make the forest structure data more accurate than those obtained in earlier studies that used TLS [
30]. Although increasing the number of scanning stations can ensure that trees are recognized as such with greater accuracy, the trade-off is that it will take more time to process the data and may lead to redundant data. In the present study, the recognition accuracy of single-station scanning was 82.56% whereas that of multi-station scanning was 97.71%—thus the accuracy of multi-station scanning was much higher. The increasing spatial resolution of laser scanners will also increase the quality of data on structural parameters of forests. Therefore, further research should focus on the development of appropriate methods to assess the quality of such data obtained using TLS.
5. Conclusions
The growth of individual trees is affected by many factors, especially in a mixed forest. Broad-leaved trees in a mixed forest of bamboo can influence the growth of bamboo and the spatial structure of the forest. In the present study, the average recognition rate of individual bamboos was 97.71%. Broad-leaved trees suppressed the growth of bamboo growing within 3 m from them but favoured the growth of bamboo when it was beyond 3 m. The competition index proposed in the present paper added canopy-related factors to the Hegyi competition index to express the competitive relationship between the trees and bamboo fully and also analysed the competitive effect of the canopy. The revised competition index indicated that large neighbours have a greater influence on the target species than small neighbours do, and the effect is also a reflection of canopy cover and lateral extrusion.
By analysing the entire 3D spatial structure obtained using TLS, we determined the degree of competition between bamboo and broad-leaved trees. These results provide a reference for future research on competition between trees in mixed forests and enhance our understanding of the spatial structure of bamboo and broad-leaved trees in mixed forests. The mix of broad-leaved trees and bamboo in the right proportions will increase the yield of bamboo and enhance the comprehensive effect of such mixed forests. Analysing the spatial structure of these forests helps to explore the patterns of growth, optimize management methods, and increase productivity. However, more accurate algorithms are needed for analysing canopy structure to detect convex and concave pockets in the crown to examine the relationship between the 3D structure of a canopy and its effect on the competition between plants for environmental resources.