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Article

Distribution Patterns of Soil Fauna in Different Forest Habitat Types of North Hebei Mountains, China

Research Center for Engineering Ecology and Nonlinear Science, North China Electric Power University, Beijing 102206, China
*
Author to whom correspondence should be addressed.
Sustainability 2022, 14(10), 5934; https://doi.org/10.3390/su14105934
Submission received: 26 March 2022 / Revised: 27 April 2022 / Accepted: 11 May 2022 / Published: 13 May 2022

Abstract

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The relationship between soil fauna distribution and forest habitat types is an ongoing concern. In this study, the distributions of soil fauna communities were investigated and compared in three forests of Betula platyphylla, Picea asperata, and Pinus sylvestris. A total of 39 groups of soil fauna belonging to four functional groups were found, with Acarina and Collembola being the dominant groups, and omnivorous and saprophagous being the dominant functional groups. An investigation on the temporal and spatial distribution of the soil fauna found similar changes in the three forests: the abundance of soil fauna was higher in August and September and lowest in May; explicit surface aggregation of the soil fauna emerged; and the density and group number decreased with the increase in soil depth. Via comparison, the total abundance of soil fauna in the B. platyphylla forest (16,772 ind m−2) was found to be higher than that in the P. asperata (12,972 ind m−2) and P. sylvestris (14,250 ind m−2) forests, and the indexes of diversity, richness and evenness of the soil fauna community in the B. platyphylla forest were the highest. Redundancy analysis showed that soil organic matter (SOC), total nitrogen (TN), and total phosphorus (TP) were positively correlated with soil fauna density, whereas pH and bulk density (BD) were negatively correlated. Compared with the two coniferous forests, the physicochemical factors positively (negatively) correlated with soil fauna density were the highest (lowest) in the B. platyphylla forest. The combined effect of these multiple factors suggests that the B. platyphylla forest recovered the most favorable conditions for the living and development of the soil fauna. The findings in this research may help us to understand the restoration effect of soil fauna in different forest habitat types, providing support for forest sustainable management in northern Hebei Mountain ecosystems.

1. Introduction

Soil fauna are an important component of terrestrial ecosystems and can exert important influences on the ecosystem structure and function [1]. By directly or indirectly participating in litter decomposition, they promote the formation of humus, accelerate the flow of soil nutrient [2,3], and regulate the basic physical and chemical properties and structure of the soil [4,5]. Due to the characterization of weak migration capability, the soil fauna is sensitive to environmental pollution or changes. Therefore, the soil fauna is often considered as an indicator of soil quality [6,7,8] or ecosystem restoration [9].
The composition and distribution of the soil fauna may greatly shift as a result of the variation in forest types [10], climatic conditions [11], altitude gradients [12], and soil factors [13]. In recent decades, the relationship between soil fauna distribution and forest habitat types has gradually become a hot topic. In the forests, generally, the under-forest microenvironment as well as the nutrient composition of litter can direct and indirectly affect the distribution patterns of the soil fauna [14,15,16]. Studies have shown that abundant soil organic matter content is related to high species richness and rich biodiversity [17]. Since the litter is the main source of soil organic matter and the plant species can greatly affect the litter quality, the vegetation community in different forests can significantly contribute to the changes in the soil fauna. Explicitly, soil community structure can be affected by an increase in tree richness [18]. Mixed stands are also likely to enhance the diversity and the richness of soil fauna [19]. Other factors associated with specific tree characteristics, such as root morphology, nutrient content, and exudates, may change after transformation, affecting the subsurface soil biota [18]. Furthermore, the interactions among species in aboveground plant communities can affect the structure and distribution of soil fauna communities, and in turn, the underground soil fauna communities can also exert a feedback influence on the composition and structure of surface plant communities [20,21]. In general, the composition of the soil fauna community is closely related to the composition of the vegetation community in the forests, and their interactions are highly complex and dynamic.
Many researchers have conducted extensive investigations on the changes in soil fauna communities and diversity in degraded ecosystems [22,23,24]. The restoration of soil fauna communities is of great significance to the structure and function of forest ecosystems during forest restoration, and soil fauna communities can be used as assessment indicators for “forest restoration benefits” [9]. In this study, we intended to investigate the distribution patterns of soil fauna in the restored forests in the North Hebei Mountains, China, in order to improve the understanding of the relationship between the forest habitat types and the soil fauna communities in the restoration process of the forests in cold temperate zone. We hypothesized that the deciduous broad-leaved forests, which promote the formation of litter and soil organic matter, can enhance the abundance and diversity of the soil fauna. By comparatively analyzing the characteristics of the soil fauna communities and the influencing factors, on the one hand, we expect to obtain the influence of distribution patterns of soil fauna by the forest habitat types, and on the other hand, to determine the environmental factors promoting and limiting the development of the soil fauna community. This research may provide references for the design of sustainable forest ecosystems in restoration.

2. Materials and Methods

2.1. Study Area

The field experiments of this study were conducted in the area of Taizicheng River catchment (40°54′50.7″–40°59′51.2″ N, 115°26′20.9″–115°28′53.3″ E) in Chongli District, Zhangjiakou, Hebei Province, China (Figure 1). The Taizicheng River catchment is situated on the transitional zone between the Inner Mongolia Plateau and North China Plain, and has a continental monsoon climate with a cold, dry winter and a five-month snow storage period during which snow can reach 1 m [25]. The summer in the catchment is warm and rainy, with an average temperature of 19 °C. The annual precipitation is 480 mm, and rainfall occurs mainly from June to September [26]. The altitude of the catchment is 1588–2134 m, and the main soil types are chestnut, meadow, brown loam, and cinnamon.
After a field investigation on the forests of the Taizicheng River catchment, three forest habitat types were selected, B. platyphylla, P. asperata, and P. sylvestris forests, to study the distribution patterns of the soil fauna communities. The selection of forest habitats was based on the following criteria: (1) the area of the habitats should be large enough; (2) the dominant species of arbor are single species; and (3) the slope and elevation should be as similar as possible. The selected three forest habitats are representative of the Taizicheng River catchment, and the natural environment characteristics of the three forests are provided in Table 1.

2.2. Collection and Processing of Samples

Soil fauna samples were collected from the three forest habitats in May, June, August, September, and November 2020. In each forest habitat, one 100 m × 100 m sample plot including three 30 m × 30 m subplots was established. Three replicate sampling points in each subplot were selected according to the diagonal method. The sampling points were required to be free of stones and human disturbances, and soil fauna nests were avoided. An 800 cm3 cutting ring (20 cm deep) was used to sample soil fauna. The sampler was driven directly into the soil, and then the soil column was excavated. This method can effectively reduce the escape of soil fauna and increase the accuracy of results. The collected soil samples were divided into four layers (0–5 cm, 5–10 cm, 10–15 cm, and 15–20 cm).
Modified Tullgren funnel extractor was used to extract the soil fauna. Eight 25 W incandescent lamps were used, and the illumination time was 48 h. All soil fauna samples were preserved in 75% alcohol. Microscopy was used for identification and counting soil fauna. Due to the limited precision of microscopes, Gastropoda, Oligochaeta, Diplopoda, Crutacea, Protura, Chilopoda, Symphyla, Arachnida, Collembola and Insecta were classified into orders or families, and some soil fauna were classified into classes such as Rotifera and Nematoda. The identification of soil fauna was based on the bibliography of the Pictorial Keys to Soil Animals of China [27] and Insect Classification Atlas [28].
To analyze the soil physical and chemical properties, including soil organic carbon (SOC), bulk density (BD), pH, total nitrogen (TN), and total phosphorus (TP), 0–5 cm soil sample was collected with 100 m3 cutting ring next to the soil fauna samples. Undisturbed soil samples collected with a ring knife were used to measure soil BD. The collected soil samples were air-dried, ground, and passed through 0.25 mm sieves for laboratory analyses. SOC was determined by the FeSO4 titration after digestion using H2SO4-K2Cr2O7 [29]. TN content was measured with an ultraviolet spectrophotometer following Kjeldahl digestion [30]. TP content was measured colorimetrically after digestion with HClO4–H2SO4 [31]. Soil pH was measured using a glass electrode at a ratio of 1:2.5 (soil/water) after shaking to equilibration for approximately 30 min [32].

2.3. Statistical Analysis

The data of soil fauna were converted into numbers per square meter (Ind/m2). Based on the classification criteria of soil fauna functional groups by Lin [33] and Zhang [34], the soil fauna was divided into four functional groups: saprophytic, predatory, omnivorous, and phytophagous. The diversity of soil fauna communities was quantified using Shannon–Wiener index (H), Margalef richness index (D), Simpson dominance index (C), and Pielou evenness index (J). The formulas of these indexes can be seen in Table 2.
Repeated one-way ANOVA measurements were performed to compare the number of groups, the density of soil fauna, community indexes, and soil properties among different forest habits, and the LSD procedure and contrasts with a probability level of 0.05 were used to identify significant differences between treatment effects. Pearson correlation analysis (two-tailed test) was used to analyze the relationship between soil fauna density and soil physicochemical factors. Statistical analyses were performed using SPSS 22.0 (IBM Corp., Armonk, NY, USA). Figures were created using the drawing software Origin 9.0 (OriginLab., Northampton, MA, USA). Redundancy analysis (RDA) was performed to analyze the relationship between soil fauna communities and environmental factors, with soil fauna communities as dependent variables and environmental factors as explanatory variables. To reduce the weight of rare species, <1% of the species were excluded before analysis.

3. Results

3.1. Soil Fauna Communities Composition

In total, 7922 soil fauna (Table 3) were collected during the entire experimental period, which belonged to 5 phyla, 12 classes, and 39 groups in the three forest habitats. The mean density of the soil fauna was 14,664 individuals/m2. Acarina and Collembola were the dominant groups, accounting for 88.2%, whereas the common groups were Diptera, Coleoptera, and Hymenoptera, accounting for 7.64%. Additionally, there were 35 rare groups, accounting for 4.16%.
According to food habits, in general, omnivorous and saprophytic soil fauna were the dominant groups in the three habitats, and a few predatory and herbivorous soil fauna were also found. Differences were observed in the composition of each functional group among the three habitats. In the B. platyphylla and P. asperata forests, the proportion of the omnivorous soil fauna was the highest, followed by the saprophagous soil fauna, whereas the proportion of the saprophagous soil fauna was higher in the P. sylvestris forest.

3.2. Distribution of Soil Fauna Communities

The soil fauna showed the lowest group number in P. sylvestris forest, and no significant difference in the soil fauna group number was found between B. platyphylla and P. asperata forests (p > 0.05). Thirty-three soil fauna groups were found in all of the three forest habitats, with Acrina and Collembola dominating. The ratios of Acarina and Collembola (A/C) in the B. platyphylla, P. asperata, and P. sylvestris forests were 0.92, 0.93, and 1.23, respectively. Thus, Collembola possessed a higher proportion than Acarina in the B. platyphylla and P. asperata forests, while in the P. sylvestris forest, the proportion of Acarina was higher. It was noticeable that the common groups of soil fauna in the three habitats differed: the B. platyphylla forest was predominated by Diptera and Coleoptera, the P. asperata forest by Hymenoptera, Diptera, and Coleoptera, and the P. sylvestris forest by Diptera and Hymenoptera. For each habitat, there were endemic groups: Elateridae, Diplatyidae, and Lithobilida were only found in B. platyphylla, P. asperata, and P. sylvestris forests, respectively. The soil fauna density in the B. platyphylla forest was significantly higher than that in the P. sylvestris and P. asperata forests (p < 0.05).
In different sampling periods, the group number and density of the soil fauna showed significant differences (p < 0.05). As shown in Figure 2, the significantly highest soil fauna group number (density) was found in August (August and September) (p < 0.05). The P. sylvestris and P. asperata forests had the highest soil fauna density in September, while B. platyphylla forests had the highest density in August. Compared with the B. platyphylla forest, the number of groups in the P. sylvestris and P. asperata forests fluctuated less. The density of soil fauna was the highest in the B. platyphylla forest in all of the months (except for May), followed by the P. sylvestris forest, and the lowest density was found in the P. asperata forest. The composition of soil fauna communities was also different between months, and some groups existed only in specific months. For example, the endemic group in June was Blattidae, and the endemic group in August was Oniscidae.
As observed from the vertical distribution, the density and group number of soil fauna in the three habitats showed clear surface aggregation (Figure 3). As soil depth increased, both the number and density of groups declined. The significance analysis showed that the group number in the 0–10 cm soil layer in the B. platyphylla forest was significantly higher than that in the 10–20 cm soil layer (p < 0.05), while in the P. asperata and P. sylvestris forests, the group number in the 0–5 cm soil layer was significantly higher than in the other soil layers.
The composition of the soil fauna community varied with different soil depth. Some groups were widely distributed in all soil layers, such as Acanthinidae, Orychomycidae, Radiatamites, and Gamomites, which were all dominant groups in the four soil layers, and Echinococcidae, Bibionidae, Hypogastruridae, Sminthuridae, and Orchesellidae, which were common groups along the soil layers. Some groups existed only in certain soil layers, for example, Schizopteridae and Corydiidae existed only in the 5–10 cm soil layer, and Oniscidae, Elateridae, Cydnidae, Stylommatophora, and Staphylinidae existed only in the 0–10 cm soil layer. There were also differences in the vertical distribution of soil fauna density among different groups. Echinococcidae and Tipulidae decreased with soil depth, whereas Stylommatophora showed an opposite trend. The densities of the Diptera larvae in each layer were roughly the same.

3.3. Diversity Characteristics

The Shannon–Wiener, richness, and evenness indexes of soil fauna communities were the highest in the B. platyphylla forest, followed by the P. asperata forest, and the lowest was found for the P. sylvestris forest. In contrast, the dominance index for the three forest habitats showed an opposite shift. Positive correlations existed between any two of the evenness, richness, and Shannon–Wiener indexes, and these three indexes were all negatively correlated with the dominance index. The diversity indexes varied temporally in the three habitats, as outlined in Figure 4. In August, the Shannon–Wiener, richness, and evenness indexes peaked, while the dominance index was at its lowest. The Shannon–Wiener index of the B. platyphylla forest was the lowest in May, while in the other months, it was higher than that of the other two habitats. The temporal variation in each diversity index was different among habitats. The Shannon–Wiener, richness, and evenness indexes of B. platyphylla were all lowest in May. For P. asperata, the lowest values of these three indexes were in November, while September exhibited the lowest values for P. sylvestris.
Different values of the diversity indexes were also observed among the soil layers (Table 4). The 0–5 cm soil layer exhibited the highest number of groups and soil fauna density among the three habitats. However, higher values of Shannon–Wiener richness and evenness indexes were found in the 5–10 cm soil layer than in the 0–5 cm layer in the B. platyphylla forest. The indexes of Shannon–Wiener and richness were highest in the 0–5 cm soil layer in the P. asperata and P. sylvestris forests. Species of soil fauna were abundant in the 5–10 cm soil layer of the B. platyphylla forest, with high uniformity and complex community structures. In the P. asperata and P. sylvestris forests, the most complex community structure was in the 0–5 cm soil layer.

3.4. Effects of Soil Physicochemical Factors on Soil Fauna Communities

Redundant analysis between the soil fauna and environmental factors was performed for two-dimensional ranking (Figure 5). The main soil physicochemical properties (SOC, SC/N, pH, TN, TP, and BD) were selected as the environmental factors to explain the community characteristics of the main groups of soil fauna. The first canonical axis was mainly determined by BD, explaining 62.64% of the total variation, and the second canonical axis mainly comprised SOC, TN, and TP, explaining 22.58% of the variation.
The results in Table 5 and Figure 5 show that soil fauna responded differently to various soil factors, and different soil fauna responded heterogeneously to the same factor. Oribatida, Actinedida, Orchesellidae, and Gamasida were more significantly related to SOC than the other groups. Oribatida, Actinedida, and Gamasida had a highly significant positive correlation with TP (p < 0.01), and a significant positive correlation with TN (p < 0.05). The C/N had a weak impact on the soil fauna. Bibionidae and Tipulidae were significantly negatively correlated with pH (p < 0.05).

4. Discussion

4.1. Spatial Distribution of Soil Fauna in Different Forest Habitats

Studies have shown that forest habitat is closely related to soil fauna density and distribution [10,39]. Through the investigation of soil fauna in the three habitats in the mountainous area of northern Hebei, we found significant differences in the composition of soil fauna communities in different habitats. The differences of the soil fauna may be largely dependent on the change in quality parameters of the soil and litter [9], which may be mainly influenced by the plant species in the habitats [21]. B. platyphylla is a type of deciduous broad-leaved tree with a rich litter that decomposes quickly, and soil fauna thrives in thick litter layers, which provide adequate habitat and foster reproduction of the soil fauna [40]. P. asperata and P. sylvestris are evergreen trees, the deciduous stage is not obvious, and the litter decomposition is slower than that of deciduous broad-leaved species [41], because the litter of P. asperata and P. sylvestris is mostly pine needles that are not easily decomposed [42,43]. Conifer litter is less rich in nutrients and less palatable for animal consumption than leaf, grass, and herb litter [44].
In this study, the P. sylvestris forest had richer shrub species, higher vegetation coverage, and higher litter layer thickness than the P. asperata forest. Moreover, plant cover, density, and litter layer thickness were positively correlated with the amount of soil fauna [16]. Thus, the density of soil fauna in the B. platyphylla forest was the highest, followed by the P. sylvestris forest, and the least density was found in the P. asperata forest. The Shannon–Wiener index, richness index, and evenness index of the soil fauna community in the B. platyphylla forest were higher than in the P. asperata and P. sylvestris forests. Although the density of soil fauna in the P. sylvestris forest was higher than that in the P. asperata forest, there were fewer groups, and the distribution of soil fauna was uneven.
Soil fauna communities are strongly correlated with soil physicochemical properties, which often influence the soil fauna in the form of multiple factors instead of a single factor [45]. Previous studies indicated that deciduous broadleaf leaves had higher nutrient contents (e.g., nitrogen and phosphorus) and specific leaf areas compared to evergreen broadleaf and conifer leaves [46]. In this study, the SOC, TN, and TP in deciduous broad-leaved forest (B. platyphylla forest) were higher than those in coniferous forest (P. asperata forest), and there was no significant difference in nutrient content (SOC, TN, TP) between the P. sylvestris forest and the B. platyphylla forest. The reason may be that there are more species and numbers of shrubs in P. sylvestris, resulting in a high degree of litter mixing. A study had showed that litter mixing can lead to an increased content of carbon, nitrogen, and phosphorous [47,48,49,50,51]. Soil fauna was negatively correlated with the soil pH and BD [52,53], as was the case in this study. Compared the three habitats, the soil physicochemical factors positively correlated with soil fauna density were the highest in the B. platyphylla forest, whereas the negatively correlated factors were the lowest in this forest. This result may explain the higher density of soil fauna in the B. platyphylla forest than the other two forests.
SOC, TN, and TP are important indicators for measuring soil fertility, which significantly increases the density of the soil fauna. The soil nutrients affect soil biota productivity via increased plant biomass production, which in turn enhances microbial growth, and thereby increases resources for soil fauna [54]. In addition, plant functional type-associated differences in litter chemistry (e.g., N and P concentration, C quantity and quality) may affect decomposability and palatability [55,56]. Previous studies had shown the biomass of Enchytraeids, Collembolans, and Mites was increased in soils with a relatively high N content [57]. Another study by Loranger-Merciris et al. [58] showed that a high content of N in leaves is attractive for soil fauna, resulting in an abundant macrofauna community. Ribeiro et al. [59] found that within a certain range, an increase in P in the soil will increase the abundance of soil nutrients and nematodes, and also change the decomposition channels of organic matter in the soil. Similar results were also found in this study. In particular, the high SOC content significantly increased the density of Oribatida, Actinedida, Orchesellidae, and Gamasida; the content of TN had a positive effect on Oribatida, Actinedida, and Gamasida; and the densities of Oribatida, Actinedida, Orchesellidae, and Gamasida increased with the content of TP.

4.2. Temporal Distribution of Soil Fauna in Different Forest Habitats

There were strong seasonal and habitat-related variations in the density and diversity of soil fauna. This result may be due to changing environmental factors [60]. The mountainous area of northern Hebei was influenced by a continental monsoon climate in East Asia, resulting in seasonal variations in temperature and precipitation [61]. Based on the results of this study, the density of soil fauna was significantly affected by the season (p < 0.05), which is consistent with the findings of Begum et al. [62]. The B. platyphylla forest was located on the shady slope of Cuiyun Mountain. In May, when we took the soil fauna samples, the soil in the B. platyphylla forest was still frozen, and the low temperatures weakened the activity of the soil fauna [63]. At the same sampling time, the P. asperata and P. sylvestris forests were located on a sunny slope, and the soil was thawed. As a result, the difference in the slope direction of the habitats led to the soil fauna communities in B. platyphylla being lower in density and group numbers than in P. asperata and P. sylvestris in May.
With an average temperature of 23 °C, August was the hottest month out of the five sampling times. It was the peak period for the reproduction and development of soil fauna due to the favorable climate and hydrothermal conditions. Compared to other months, soil fauna’s Shannon–Wiener index, richness index, and evenness index were higher this month. There were more litter accumulations in September. Therefore, soil fauna groups tended to be denser and more numerous in August and September. Although the soil fauna density was the highest in September in the P. sylvestris forest, the diversity index was conversely the lowest, mainly due to the conditions in this forest being beneficial for the growth of a few groups of soil fauna which had a higher abundance. In November, the density of soil fauna was higher in all three habitats, but the number of groups was lower because low temperatures made most of the soil fauna groups unable to survive, and thus the number of species of soil fauna declined. Nevertheless, the accumulation of litter in November increased the density of those groups that feed on litter, such as Acarina. Thus, the total density of soil fauna remained high in November.

5. Conclusions

The same dominant groups of the three forests were found, namely, Acarina and Collembola, which demonstrated universal adaptability to different microhabitats. Under the influence of the dominant groups, omnivorous and saprophagous were the main functional groups. The temporal and spatial distributions of soil fauna in the three habitats were similar. The abundance of soil fauna exhibited pronounced seasonal variations over five months and was higher in August and September, due to the favorable climate and hydrothermal conditions. Meanwhile, owing to air permeability and food resources, the majority of soil fauna lived in the topsoil. In the comparison, the density and group number of soil fauna in the B. platyphylla forest were found to be the highest, and the Shannon–Wiener, richness, and evenness indexes of soil fauna communities were the highest in the B. platyphylla forest, followed by the P. asperata forest, and the lowest was found in the P. sylvestris forest, which indicated that the stability and diversity of soil fauna communities in B. platyphylla forest were higher.
Our results suggest that the abundance and diversity of the soil fauna community were higher in the deciduous broad-leaved forest (B. platyphylla forest) than in the coniferous forests (P. asperata and P. sylvestris forests), which was the result of the combined effects of SOC, TP, TN, pH, and BD, among which the soil physicochemical factors positively correlated with soil fauna density were the highest in the deciduous broad-leaved forest, whereas the negatively correlated factors were the lowest in this forest. Therefore, the deciduous broad-leaved forest (B. platyphylla forest) is more suitable for the protection of biodiversity during forest restoration, and this study is provides a reference for the forest restoration of the northern Hebei Mountains.

Author Contributions

Conceptualization, H.Z. and T.H.; methodology, H.Z. and Q.L.; software, Q.L.; writing—original draft preparation, H.Z., Q.L. and T.H.; writing—review and editing, T.H. and Q.L.; visualization, Q.L.; supervision, H.Z. and T.H.; funding acquisition, H.Z.; data curation, Y.F. and S.Z. All authors have read and agreed to the published version of the manuscript.

Funding

This research was supported by the National Science and Technology Major Project for Water Pollution Control and Treatment (2017ZX07101002).

Institutional Review Board Statement

Not applicable.

Informed Consent Statement

Not applicable.

Data Availability Statement

Sources of all the soil fauna data used in this study are described in Section 2.2. The data are shown in the figures and tables of the paper and are available upon request.

Conflicts of Interest

The authors declare no conflict of interest.

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Figure 1. Location of the study site, which is located in Chongli District, Zhangjiakou, Hebei Province, China. The black marking point is the Taizicheng River catchment.
Figure 1. Location of the study site, which is located in Chongli District, Zhangjiakou, Hebei Province, China. The black marking point is the Taizicheng River catchment.
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Figure 2. The dynamics of individual density and group number of soil fauna in different habitats. Capital letters indicate temporal differences within habitats at the p < 0.05 level, while lowercase letters indicate spatial differences within seasons at the p < 0.05 level.
Figure 2. The dynamics of individual density and group number of soil fauna in different habitats. Capital letters indicate temporal differences within habitats at the p < 0.05 level, while lowercase letters indicate spatial differences within seasons at the p < 0.05 level.
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Figure 3. Vertical distribution of (a) individual density and (b) group number of soil fauna in different habitats. Capital letters indicate soil layer differences within habitats at the p < 0.05 level, while lowercase letters indicate spatial differences within soil layer at the p < 0.05 level.
Figure 3. Vertical distribution of (a) individual density and (b) group number of soil fauna in different habitats. Capital letters indicate soil layer differences within habitats at the p < 0.05 level, while lowercase letters indicate spatial differences within soil layer at the p < 0.05 level.
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Figure 4. (a) Shannon–Wiener index, (b) Margalef richness index, (c) Pielou evenness index, and (d) Simpson dominance index of the soil fauna in different habitats. BH: Betula platyphylla; YS: Picea asperata; ZZS: Pinus sylvestris. Capital letters indicate spatial differences within seasons at the p < 0.05 level, while lowercase letters indicate temporal differences within habitats at the p < 0.05 level.
Figure 4. (a) Shannon–Wiener index, (b) Margalef richness index, (c) Pielou evenness index, and (d) Simpson dominance index of the soil fauna in different habitats. BH: Betula platyphylla; YS: Picea asperata; ZZS: Pinus sylvestris. Capital letters indicate spatial differences within seasons at the p < 0.05 level, while lowercase letters indicate temporal differences within habitats at the p < 0.05 level.
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Figure 5. Results of redundancy analysis for the main groups of the soil fauna in association with selected properties of soil. Red arrows represent soil properties labeled as: soil organic matter (SOC), soil TC/TN (C/N), total nitrogen (TN), total phosphorus (TP), bulk density (BD); blue arrows represent groups of soil fauna labeled as: Tipulidae (Tip), Isotomidae (Iso), Echinococcidae (Ech), Oribatida (Ori), Bibionidae (Bib), Actinedida (Act), Hypogastruridae (Hyp), Diptera (Dip), larvae (Lar), Formicidae (For), Sminthuridae (Smi), Orchesellidae (Orc), Gamasida (Gam).
Figure 5. Results of redundancy analysis for the main groups of the soil fauna in association with selected properties of soil. Red arrows represent soil properties labeled as: soil organic matter (SOC), soil TC/TN (C/N), total nitrogen (TN), total phosphorus (TP), bulk density (BD); blue arrows represent groups of soil fauna labeled as: Tipulidae (Tip), Isotomidae (Iso), Echinococcidae (Ech), Oribatida (Ori), Bibionidae (Bib), Actinedida (Act), Hypogastruridae (Hyp), Diptera (Dip), larvae (Lar), Formicidae (For), Sminthuridae (Smi), Orchesellidae (Orc), Gamasida (Gam).
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Table 1. Natural environment characteristics of the three forest habitats.
Table 1. Natural environment characteristics of the three forest habitats.
Forest TypeAltitude
(m)
Latitude and LongitudeAspectDominant SpeciesTree LayerBush Layer
Betula platyphylla (BH)1948115°26′36″ E
40°58′48″ N
Northeast directionBetula platyphyllaBetula platyphylla
Malus baccata
Rosa bella
Rubus swinhoei
Abelia biflora
Crataegus pinnatifida
Salix cheilophila
Picea asperata (YS)1890115°27′40″ E
40°57′43″ N
Southwest directionPicea asperataPicea asperata
Armeniaca sibirica
Malus baccata
Potentilla fruticose
Rosa bella
Spiraea salicifolia
Pinus sylvestris (ZZS)1900115°27′40″ E
40°57′36″ N
Southwest directionPinus sylvestrisPinus sylvestris
Betula platyphylla
Larix gmelinii
Malus baccata
Potentilla fruticose
Rosa bella
Rubus swinhoei
Abelia biflora
Salix cheilophila
Spiraea salicifolia
Table 2. The formulas of diversity indexes.
Table 2. The formulas of diversity indexes.
Diversity IndexesFormulas
Shannon–Wiener index [35] H = ( n i / N ) ln ( n i / N )
Margalef richness index [36] D = ( S 1 ) / ln N
Simpson dominance index [37] C = ( n i / N ) 2
Pielou evenness index [38] J = H / ln S
ni is the number of individuals in the group; N is the total number of individuals in all groups; S is the number of groups.
Table 3. Mean density (Ind/m2) and percentage (%) of soil fauna in different habitats.
Table 3. Mean density (Ind/m2) and percentage (%) of soil fauna in different habitats.
BHYSZZSFg.
Ind/m2%DegInd/m2%DegInd/m2%Deg
Monoptera33.33 0.20%+50.00 0.39%+22.22 0.16%+S
Carabidae44.44 0.26%+16.67 0.13%+5.56 0.04%+Pr
Oniscidae22.22 0.13%+11.11 0.09%+5.56 0.04%+O
Tipulidae211.11 1.26%++155.56 1.20%++111.11 0.78%+S
Geophilidae116.67 0.70%+38.89 0.30%+33.33 0.23%+Pr
Isotomidae1733.33 10.33%+++1133.33 8.74%++1016.67 7.13%++O
Eosentomidae33.33 0.20%+66.67 0.51%+33.33 0.23%+S
Phlaeothripidae150.00 0.89%+33.33 0.26%+38.89 0.27%+O
Echinococcidae3161.11 18.85%+++2444.44 18.84%+++2500.00 17.54%+++O
Oribatida3088.89 18.42%+++2155.56 16.62%+++3144.44 22.07%+++S
Elateridae16.67 0.10%+0.00 0.00% 0.00 0.00% Ph
Juliformia38.89 0.23%+5.56 0.04%+5.56 0.04%+S
Schizopteridae11.11 0.07%+11.11 0.09%+0.00 0.00% Ph
Bibionidae250.00 1.49%++166.67 1.28%++127.78 0.90%+Pr
Actinedida1794.44 10.70%+++1538.89 11.86%+++2072.22 14.54%+++S
Diplatyidae0.00 0.00% 5.56 0.04%+0.00 0.00% O
Coleoptera larvae144.44 0.86%+66.67 0.51%+44.44 0.31%+Ph
Hypogastruridae961.11 5.73%++1066.67 8.22%++966.67 6.78%++O
Diptera larvae322.22 1.92%++161.11 1.24%++166.67 1.17%++Ph
Enchytraeidae27.78 0.17%+22.22 0.17%+22.22 0.16%+S
Lithobilidae0.00 0.00% 0.00 0.00% 11.11 0.08%+Pr
Cydnidae27.78 0.17%+11.11 0.09%+0.00 0.00% Ph
Stylommatophora22.22 0.13%+0.00 0.00% 5.56 0.04%+S
Curculionidae100.00 0.60%+16.67 0.13%+27.78 0.19%+Ph
Araneae61.11 0.36%+33.33 0.26%+38.89 0.27%+Ph
Mycetophilidae166.67 0.99%+83.33 0.64%+44.44 0.31%+S
Scorpionida44.44 0.26%+77.78 0.60%+33.33 0.23%+S
Elateridae22.22 0.13%+11.11 0.09%+5.56 0.04%+Ph
Formicidae83.33 0.50%+166.67 1.28%++194.44 1.36%++O
Staphylinidae61.11 0.36%+44.44 0.34%+5.56 0.04%+Pr
Muscidae138.89 0.83%+116.67 0.90%+61.11 0.43%+S
Sminthuridae922.22 5.50%++927.78 7.15%++766.67 5.38%++O
Orchesellidae711.11 4.24%++327.78 2.53%++627.78 4.41%++O
Lumbricidae22.22 0.13%+16.67 0.13%+5.56 0.04%+S
Gamasida1994.44 11.89%+++1777.78 13.70%+++1983.33 13.92%+++S
Katydidae27.78 0.17%+27.78 0.21%+5.56 0.04%+Ph
Rotifera100.00 0.60%+66.67 0.51%+77.78 0.55%+O
Nematoda100.00 0.60%+105.56 0.81%+38.89 0.27%+O
Corydiidae5.56 0.03%+11.11 0.09%+0.00 0.00% O
Density16,772 12,972 14,250
Group number37 36 34
BH: Betula platyphylla; YS: Picea asperata; ZZS: Pinus sylvestris (The meaning of BH, YS, and ZZS is the same in the following tables and figures). Deg.: degree of dominancy, +++ indicates a dominant group, ++ indicates a common group, and + indicates a rare group. Fg.: functional group, with O indicating omnivores, S indicating saprozoics, Pr indicating predators, and Ph indicating phytophages.
Table 4. Diversity indexes of soil fauna in different soil layers.
Table 4. Diversity indexes of soil fauna in different soil layers.
SampleSoil Layer
(cm)
Diversity
(H)
Abundance
(D)
Evenness
(J)
Dominance
(C)
BH0–52.4974.8720.6970.115
5–102.5425.0570.7150.114
10–152.4534.1310.7440.123
15–202.4403.4540.8010.115
YS0–52.4904.5230.9300.343
5–102.4854.4460.7310.113
10–152.3093.4370.7470.128
15–202.2553.2390.7660.143
ZZS0–52.3054.4070.6650.132
5–102.3013.8170.7060.133
10–152.2333.2590.7330.142
15–202.1052.6790.7590.153
Table 5. Pearson correlation matrix between main groups of soil fauna and quality parameters of soil.
Table 5. Pearson correlation matrix between main groups of soil fauna and quality parameters of soil.
pHSOCTNTPBDC/N
Tip−0.720 *0.3590.260.159−0.30.26
Iso−0.4730.3360.1360.172−0.4850.638
Ech−0.4810.4730.2710.368−0.5590.627
Ori−0.380.818 **0.737 *0.861 **−0.5780.143
Bib−0.700 *0.4110.2930.226−0.3620.313
Act−0.2690.732 *0.763 *0.832 **−0.316−0.251
Hyp−0.356−0.0690.056−0.0960.461−0.477
Dip−0.5710.5450.380.335−0.5060.478
For−0.0970.0920.3150.2280.471−0.844 **
Smi−0.4330.009−0.051−0.1450.080.197
Orc−0.4650.774 *0.6580.787 *−0.5740.266
Gam−0.4850.761 *0.694 *0.812 **−0.3950.085
Note: * p < 0.05; ** p < 0.01.
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Zhang, H.; Lin, Q.; Huang, T.; Feng, Y.; Zhang, S. Distribution Patterns of Soil Fauna in Different Forest Habitat Types of North Hebei Mountains, China. Sustainability 2022, 14, 5934. https://doi.org/10.3390/su14105934

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Zhang H, Lin Q, Huang T, Feng Y, Zhang S. Distribution Patterns of Soil Fauna in Different Forest Habitat Types of North Hebei Mountains, China. Sustainability. 2022; 14(10):5934. https://doi.org/10.3390/su14105934

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Zhang, Huayong, Qingxia Lin, Tousheng Huang, Yu Feng, and Shijia Zhang. 2022. "Distribution Patterns of Soil Fauna in Different Forest Habitat Types of North Hebei Mountains, China" Sustainability 14, no. 10: 5934. https://doi.org/10.3390/su14105934

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