Abstract
Aims
Non-native trees are widely used to prevent soil erosion in the Loess Plateau. We aim to investigate the effects of non-native tree plantations on the communities of understory plants and soil macroinvertebrates.
Methods
The soil, understory plants and soil macroinvertebrates were sampled in non-native Robinia pseudoacacia (RP) and Populus ×canadensis (PC) stands, and native Armeniaca sibirica (AS) stands.
Results
Abundance and richness of understory plants were significantly greatest in the RP stands. The macroinvertebrates collected belonged to 13 orders across all stands. At the community level, richness did not differ among the stand types, but the abundance was significantly greatest in the PC stands. At the functional group level, phytophages and detritivores preferred the PC stands, whereas predators preferred the RP stands. The generalist macroinvertebrate abundance was significantly greatest in the PC stands. Moreover, the community composition of understory plants clearly differed among the stand types; that of soil macroinvertebrates in the PC stands was separated from the other stand types, which was mediated by environmental variables, especially the soil temperature and soil organic carbon.
Conclusions
Non-native tree plantations had positive effects on the overall abundance of understory plants and soil macroinvertebrates, but their effects on functional groups were inconsistent, associated with tree identity. The selection of non-native tree should be considered in term of multiple trophic levels, if afforestation has biodiversity objectives.
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Introduction
Ecosystem degradation may be induced by intensive human activities and climatic changes, and can result in ecosystem function decline and ecosystem service loss (Jackson and Hobbs 2009; Allan et al. 2015). Afforestation is one of the most common management approaches to enhance ecosystem characteristics and restore degraded environments (Pimentel and Kounang 1998). Due to their rapid growth rates, non-native tree species have been widely planted to restore ecosystems worldwide (Zobel et al. 1987; Degomez and Wagner 2001).
Although non-native tree species can restore degraded ecosystems faster than native plants, they may have different effects on the physical environment when they are planted within local ecosystems (Qiu et al. 2010; Orozco-Aceves et al. 2015; Kou et al. 2016), resulting in changes to the composition, structure, and function of local ecosystems (Irwin et al. 2014; Horák et al. 2019). Plants are primary producers and provide consumers with both food and habitat. Plantations with different species may affect resource availability and habitat properties, and have cascading effects on local fauna (Bertheau et al. 2009; Sylvain and Buddle 2010; Salamon et al. 2011). Despite the well-documented impacts of non-native tree plantations (e.g., Robinia pseudoacacia, Pinus radiate, P. contorta) on the composition (Meers et al. 2010; Kasel et al. 2015), structure and cover of understory plants (Roberge and Stenbacka 2014; Huang et al. 2019) and the recognized links between aboveground and belowground biotic communities (Wardle et al. 2004; Kayler et al. 2010; Kivlin and Hawkes 2011), our understanding of the cascade effects of non-native tree plantations on soil fauna is poor (Sylvain and Buddle 2010; Roberge and Stenbacka 2014).
Soil arthropods are a common, abundant, and functionally important group of organisms in most ecosystems (Bardgett and van der Putten 2014; Wagg et al. 2014). Their activities have significant impacts on soil properties and ecosystem processes and functions (Wagg et al. 2014), such as decomposition and nutrient cycling (Bardgett and van der Putten 2014). Understanding the responses of soil arthropods to non-native tree plantations can inform efforts to conserve biodiversity and ecological processes. Some studies have examined the effects of non-native plants on the abundance, diversity, and composition of ground and belowground arthropods, and the reported effects appear to be idiosyncratic (e.g., Sax 2002; Brockerhoff et al. 2008; Quine and Humphrey 2010). Such differences reveal that the responses of arthropods are associated with non-native tree species identity, as well as the functional group to which they belong. To date, most studies only examined a single non-native species, and focused on a single or very few response variables within one ecosystem (Hulme et al. 2013). To better understand the consequences of non-native species plantation, studies are needed that include a greater variety of non-native plant species and response variables—such as soil properties, understory plant communities, invertebrate predators, phytophages, detritivores, and omnivores—within one ecosystem. Such data are valuable as they provide a more comprehensive assessment of the impacts of non-native plants on the local ecosystems (Lazzaro et al. 2017).
The Loess Plateau of China, one of the most severely eroded regions of the world, has received substantial research attention in China (Jiang et al. 2003; Deng et al. 2012; Kou et al. 2016; Yang et al. 2019). The destruction of vegetation cover, as a result of long-term human activities, has led to severe soil erosion (Chen et al. 2015). Afforestation has been widely implemented, especially under the Grain for Green program, which increased vegetation coverage on the Loess Plateau from 31.6% in 1999 to 59.6% in 2013 (Chen et al. 2015). Non-native Robinia pseudoacacia (RP) and Populus ×canadensis (PC) have been extensively planted because of their rapid growth rates (Yang et al. 2006). The impacts of plantations on soil properties (e.g., Qiu et al. 2010; Xu et al. 2014) and native plant communities (e.g., Jiang et al. 2003; Chen and Cao 2014) have been well-documented for the Loess Plateau. There is a lack of studies, however, on how soil macroinvertebrates—as a major component of many ecosystems—are affected by non-native tree plantations as well as associated understory plant communities and soil properties that may underlie macroinvertebrate responses. This knowledge gap may preclude us from determining whether or not non-native species should be widely planted in degraded ecosystems.
The aim of our work was to assess the impacts of two non-native plantations (RP and PC) on soil properties, diversity of understory plants, and soil macroinvertebrates and compare these with a native Armeniaca sibirica (AS) ecosystem. Specifically, we answer the following questions: (1) Do non-native tree plantations influence soil properties in comparison with the native tree plantation? (2) Do non-native plantations alter community composition and structure of understory plants and macroinvertebrates in comparison with the native tree plantation? (3) Are any differences in macroinvertebrate communities associated with plantation-induced changes in soil properties and understory plant communities?
Materials and methods
Study site
We selected the Yangjuangou Catchment (36°42′ N, 109°31′ E; Fig. S1), which is located in the middle region of the Loess Plateau near Yan’an, Shaanxi, China, as the study area. The region has a semi-arid continental climate with an average annual rainfall of 535 mm, and the rainfall is concentrated mainly between July and September and varies greatly from year to year (Wang et al. 2011). The soil is mainly derived from loess, with a texture that ranges from fine silt to silt, and is vulnerable to erosion (Fu et al. 2000). Long-term human activities (i.e., cultivation) have destroyed most natural vegetation, and after approximately 20 years of comprehensive management, soil erosion has been significantly mitigated. Robinia pseudoacacia (RP) is a broad-leaved and nitrogen fixing tree, Populus ×canadensis (PC) is a deciduous tree, and both of them are fast-growing tree. Armeniaca sibirica (AS) is one of the cash forest species in arid and semi-arid areas. All three tree species have been widely used in restoration projects at the study site, especially in “Grain for Green” projects by the Chinese government. This is a large-scale project that requires farmers to reserve a part of their sloping farmland for trees, shrubs, or grasses (Zhou et al. 2009).
Sampling design
We separately selected three stands (1–2 ha; >700 m apart) with similar topography and plantation ages (15 yr) for each tree species. The RP, PC and AS were planted on abandoned arable fields, where corn was planted previously. The soil in each stand was similar prior to planting and subsequent differences in soil properties reflect the effects of different tree species. Three plots (10 × 10 m) about 50 m apart were established in each stand, and the distance from the plots to the stand edge is more than 30 m. The understory plants diversity was investigated, and soil and macroinvertebrates were sampled in four 1 m × 1 m subplots of each plot from the 22 th June to 14 th July 2018 with similar weather conditions. There was no rainfall during this period.
Leaf litter (50 × 50 cm) was collected and one quadrat (50 × 50 × 30 cm depth) was excavated in each subplot; any organisms were recovered by hand sorting. The collected macroinvertebrates were preserved in a 70% alcohol solution in the field, and brought back to the laboratory for identification. In the laboratory, we counted the macroinvertebrates under a dissecting microscope. The order and family was identified for all taxa based on Yin (2001) and Zheng and Gui (2004).
Estimation of plants and soil parameters
We investigated the herbaceous vegetation to determine plant density (individuals/m2), richness, and vegetation cover (%) in each subplot. Soil cores were taken at each subplot. We measured the soil bulk density, water content, soil organic carbon, total nitrogen, and soil pH. Soil bulk density and water content were determined gravimetrically by drying the soil samples at 105 °C for 48 h. Soil organic carbon (SOC) was measured using K2Cr2O7–H2SO4 following the Walkley–Black oxidation method, and soil total N was determined using an elemental analyzer (vario EL III, Elementar Analysensysteme GmbH, Langenselbold, Germany). Soil pH was measured in a soil-water suspension (1:5 soil-water ratio) with pH analyzer (FE20, METTLER TOLEDO, Shanghai, China). Soil temperature was measured with a digital thermometer at the depth of 5, 10 and 20 cm in each subplot during soil macroinvertebrate sampling.
Data analysis
The macroinvertebrate data from the litter and soil were combined, and the results were converted into individuals per square meter (ind. m−2). We used linear mixed effects models (function lme in the package nlme of R software; Bates et al. 2015) to examine the effects of stand type (Robinia pseudoacacia, Populus ×canadensis and Armeniaca sibirica) on the soil properties, understory plants, and soil macroinvertebrates (including the overall macroinvertebrates, main taxonomic groups, functional groups, and the groups of forest generalist and specialist species). The stand type was fitted as a fixed effect, and nested within plots, which were modelled as random effects. Each response variable (soil properties, understory plants, and soil macroinvertebrates) was log-transformed where necessary to meet the assumptions of parametric tests. If significant effects were detected, we used post hoc Tukey tests (multcomp package of R software; Hothorn et al. 2008) to evaluate significant differences between stand types. All the statistical tests were evaluated using α = 5%.
Ordination of multidimensional data on community composition is an important multivariate statistical method widely used in the biodiversity research. Principal component analysis (PCA) and detrended correspondence analysis (DCA) are unconstrained ordination method, and canonical correspondence analysis (CCA) is a constrained ordination method. PCA was used to evaluate the effects of stand type on community composition of macroinvertebrates and understory plants. DCA was conducted to estimate the length of the gradient on the log-transformed macroinvertebrate abundance data. Due to the long gradient length (> 4 SD), CCA was chosen to determine the relative contributions of measured environmental parameters (soil and understory plant) to the community composition of macroinvertebrates with a Monte Carlo test, and stand was included as a covariate. All statistical analyses were performed in R 3.5.1 and CANOCO v5 (ter Braak and Šmilauer 2012).
Results
Soil properties
Soil properties differed among the non-native and native plantations (Table 1). The water content, soil temperature, and soil pH significantly differed among the stand types, with the highest values being observed in the PC stands; no differences were observed for bulk density, SOC, and TN (Table 1).
Macroinvertebrates
Abundance, richness, and community composition
The macroinvertebrate abundance within specific taxonomic groups significantly differed among the non-native and native plantations (Table 2; Fig. 1). The macroinvertebrates collected belonged to 13 orders (i.e., Tubificida, Araneae, Polydesmida, Scutigeromorpha, Scolopendromorpha, Geophilomorpha, Protura, Dermaptera, Hemiptera, Coleoptera, Lepidoptera, Diptera, and Hymenoptera) and 25 families (Table S1). Coleoptera (33.33%), Araneae (13.33%), Polydesmida (12.73%), Geophilomorpha (9.70%), and Hemiptera (7.27%) were the five most abundant orders. Of them, three groups significantly differed among the stand types (Table 2; Fig. 1). The abundances of Geophilomorpha and Coleoptera were greatest in the PC stands, whereas that of Araneae was greatest in the RP stands (Table 2; Fig. 1).
At the community level, macroinvertebrate abundance was significantly greater in the PC stands than in the RP and AS stands (Table 2; Fig. 2), whereas the richness did not differ among the stand types (Table 2; Fig. 2). The PCA results showed that community composition varied among the stand types (Fig. 3), and the first and second axes explained 28.9% and 43.3% of the total variation, respectively. Paradoxosomatidae and larval Scarabaeidae were the main groups associated with the separation of PC1 axis; Liocranidae, Scolopendridae and Geophilidae were the main groups associated with the separation of PC2 axis. Larval Scarabaeidae was most associated with the PC stands, and Tingidae was most associated with the RP and AS stands.
Functional groups and forest generalist and specialist groups
The macroinvertebrate abundance and richness within specific functional groups significantly differed among the non-native and native plantations (Fig. 4). The abundance of predators and omnivores did not significantly differ among the stand types (Table 2; Fig. 4a, d), whereas that of phytophages was significantly more abundant in the PC stands than in the RP and AS stands (Table 2; Fig. 4b), and detritivores were significantly more abundant in the PC stands than in the RP stands (Table 2; Fig. 4c). Predator richness in the RP stands was significantly greater than in the PC stands (Table 2; Fig. 4e), and the detritivore richness in the PC stands was significantly greater than in the RP and AS stands (Table 2; Fig. 4g); the richness of phytophages and omnivores did not significantly differ among the stand types (Table 2; Fig. 4f, h).
The abundance of forest generalist species significantly differed among stand types, with their abundance in the PC stands being greater than in the AS stands, whereas the forest specialist abundance, and the generalist and specialist richness did not differ among stand types (Table 2).
Understory plants
For all stands, the most represented plant families were Asteraceae, Rosaceae, and Rubiaceae (Table S2). Most of the understory plants (except Parthenium hysterophorus) were native to the Loess Plateau. Plant abundance (F = 15.43, P < 0.001, LMMs; Fig. 2) and coverage (F = 31.82, P < 0.001, LMMs) were significantly greater in the RP stands than in the PC and AS stands. The plant richness in the RP and PC stands was twice as high as in the AS stands (F = 14.49, P < 0.001, LMMs; Fig. 2). The PCA results showed that plant community composition varied among the stand types (Fig. 5), and the first and second axes explained 27.7% and 48.0% of the total variation, respectively. Rubia sp., Artemisia annua and Agropyron cristatum were the main species associated with the separation of PC1 axis; Rubus coreanus, Saussurea sp., Viola betonicifolia and Parthenium hysterophorus were the main species associated with the separation of PC2 axis. Viola betonicifolia and P. hysterophorus were most associated with the RP stands; A. annua and Rubia sp. were most associated with the PC and AS stands, respectively.
Contributions of environmental variables to community composition of macroinvertebrates
CCA showed that first and second axes explained 13.45% and 24.53% of the data, respectively (Fig. 6). CCA also showed that different environmental variables had different influences on macroinvertebrate community composition (Fig. 6). The effects of soil temperature (F = 2.3; P = 0.014) and SOC (F = 2.2; P = 0.016) on macroinvertebrate communities were significant as shown under the Monte Carlo permutation test, whereas the remaining variables were not shown to be significant with the same test (plant coverage, P = 0.124; TN, P = 0.36; pH, P = 0.386; plant density, P = 0.69; bulk density, P = 0.84; plant richness, P = 0.864; water content, P = 0.7).
Discussion
Effects of stand type on soil macroinvertebrates and understory plants
There has been considerable debate regarding the impact of non-native plantations on soil macroinvertebrates (Bonham et al. 2002; Robson et al. 2009; Quine and Humphrey 2010). In general, non-native plantations are popularly regarded as “biological deserts” (Allen et al., 1995a), and seen to have negative effects on invertebrate communities compared with native plantations (Brockerhoff et al. 2008). However, some studies showed that non-native plantations have similar (Sax 2002) or higher (Quine and Humphrey 2010) diversity of invertebrates than native plantations. For example, Quine and Humphrey (2010) reported that the richness of all sampled invertebrate groups was higher in non-native Norway spruce (Picea abies) stands compared to native oak (Quercus robur) stands. In this study, the stand type had no significant effect on macroinvertebrate richness, but did affect their overall abundance. The two non-native tree plantations had different effects on the macroinvertebrate abundance than the native trees. Overall macroinvertebrate abundance was significantly greater in the non-native PC stands than in the native AS stands, but no significant difference was observed between the non-native RP and AS stands. The result implied that the impacts of non-native plantations on soil macroinvertebrates depend on species identity. Therefore, studies on the impacts of non-native plantations on the soil macroinvertebrates should focus on various non-native species rather than one, which could contribute to more in-depth understanding of the consequences of non-native plantations within an ecosystem.
Native and non-native plantations may have different effects on specific macroinvertebrate taxa. Our results showed that different orders had contrasting responses to stand type; Polydesmida and Hemiptera abundances showed no significant responses to stand type; Geophilomorpha and Coleoptera abundances were greatest in the PC stands, whereas Araneae abundances were greatest in the RP stands. This result is in line with several studies (Gerber et al. 2008; Gallé et al. 2015), which suggested that non-native plantations can have different effects on local macroinvertebrate assemblages. Such differences may be related to the microhabitat preferences of different taxa. For example, the higher abundance of Geophilomorpha and Coleoptera in PC stands may be related to the higher water content, as it is known that Geophilomorpha (Dong et al. 2014) and Coleoptera (Liu et al. 2013) prefer sites with these conditions.
The effects of non-native and native plantations on understory plant diversity are under debate (Brockerhoff et al. 2008; Bremer and Farley 2010; Thijs et al. 2014; Horák et al. 2019). While some researches showed that native plantations perform better for the biodiversity restoration than the non-native plantations (Hartley 2002; Brockerhoff et al. 2008), others studies suggested that non-native plantations are not always negative (Brockerhoff et al. 2008; Bremer and Farley 2010; Horák et al. 2019). Peloquin and Hiebert (1999) reported that RP significantly reduced the natural diversity of herbaceous plants in Quercus velutina savannas and woodland dune communities in northern Indiana. Likewise, Boothroyd-Roberts et al. (2013) reported that hybrid poplar plantations contributed to the understory plant richness in comparison with adjacent abandoned fields. However, Sitzia et al. (2012) did not record a significant difference in the richness and diversity of the understory plant community between 32 RP stands and paired native stands in the Eastern Alps. In this study, we found that understory plant richness was significantly greater in the RP and PC stands than in the AS stands, and the understory plant abundance was significantly greater in the RP stands than in the PC and AS stands. This indicated that non-native plantations had positive effects on the understory plant diversity in our study site. In addition, we also found that most of the understory plants in all plantations were native to Loess Plateau, despite the fact that non-native plantations often favored exotic plant species (Allen et al. 1995b; Ito et al. 2004; Paritsis and Aizen 2008). In combination with previous studies, the results indicate that the effects of non-native plantations on the diversity of understory plants are ambiguous.
Effects of stand type on macroinvertebrate functional groups
While there is a growing body of evidence that non-native plants can influence soil fauna, their impact on different invertebrate feeding guilds has received relatively less attention (Gratton and Denno 2005; Gerber et al. 2008). Gerber et al. (2008) pointed out that studies addressing multiple trophic levels, e.g., predators, phytophages, detritivores, and omnivores, may yield additional insight into how non-native species alter ecosystem patterns and processes. They reported that the abundances of predators, phytophages, and detritivores were all significantly greater in non-native Fallopia stands than in native grassland and bush stands (Gerber et al. 2008). In this study, we found that the distribution patterns of each functional group were different among the stand types. Firstly, the abundance and richness of phytophages and detritivores were greatest in the PC stands, albeit only significantly for phytophage abundance. The PC stands had more ground litter and decaying vegetation (Zhu, pers. obs.), which may provide alternative food resources and preferred microclimate conditions for detritivores (Litt et al. 2014). Indeed, in the review by Litt et al. (2014), the authors also showed that detritivores were most likely to benefit from non-native plants. Secondly, predator abundance and richness were greatest in the RP stands. This pattern may be associated with the greater understory plant abundance (Dassou and Tixier 2016) and coverage in the RP stands than in the other two stands; the greater plant abundance and coverage may provide alternative food and suitable microhabitat for the insects on herbaceous plants, and therefore have a positive effect on the predators through bottom-up effects. Finally, omnivore abundance and richness did not significantly vary among the three stand types. This finding is consistent with the work of Oxbrough et al. (2016), who reported that the species richness of omnivores was similar between the stands of non-native Norway spruce (Picea abies) and native ash (Fraxinus excelsior). This result suggests that omnivores may be well represented in all stands because they can feed on a greater diversity of food types than other functional groups.
Effects of environmental variables on soil macroinvertebrates
Bardgett and van der Putten (2014) showed that spatial patterns in soil biota at local scales were often explained by variation in the physical and chemical properties of the soil. Soil fauna have low migration ability and small home ranges and, therefore, any changes in their microenvironment, such as soil properties or understory plant communities, could have a significant effect on their communities. The characteristics of plant communities (such as species richness and cover) and soil properties (such as soil temperature, water content, bulk density, organic matter, and pH) are often correlated with soil fauna composition (Cole et al. 2008; Huerta and Wal 2012; Wu et al. 2015). In this study, we also found that observed differences in community structure among the three stand types were mediated by environmental variables; soil temperature was the most important factor determining macroinvertebrate composition, as it explained the greatest proportion of the total variation. It is generally accepted that soil temperature is closely associated with macroinvertebrate community composition (Perner and Malt 2003; Liu et al. 2015), and has a positive effect on their abundance (Liu et al. 2015; Robinson et al. 2018), because soil temperature can affect feeding activity (Gongalsky et al. 2008) and egg laying of soil fauna (Willis et al. 2008).
Beside the soil temperature, the water content and soil pH also significantly differed among the stand type, with both of them being greater in the RP and PC stands than in the AS stands. These differences may be potential reasons which lead to the relatively higher macroinvertebrate abundance in the RP and PC stands, because the water content (Wu et al. 2015) and soil pH (Oxbrough et al. 2010) have positive effects on the invertebrate abundance. At the functional group level, the greater abundance and richness of phytophages and detritivores in the PC stands may also be associated with the higher water content, because Scarabaeidae larvae and Oligochaeta, which are the dominant phytophages and detritivores, respectively, prefer soil with higher water content. Additionally, RP is a nitrogen fixing and therefore may be expected to impact soil nitrogen. However, despite TN being greatest in the RP stands, there were no significant differences among stands. This may be attributed to the young age of the RP forests (i.e., 15 yr) in this study, because the TN may increase along the chronosequence of 10, 20, 30, and 40 yr RP plantations in the Loess Plateau (Kou et al. 2016).
It has been proposed that the variation in plant communities seen following the introduction of non-native plants may alter resource availability and habitat properties and may, therefore, have cascading effects on local fauna (Robson et al. 2009; Litt et al. 2014). For example, the higher species richness of invertebrates in the eucalypt woodland than pine plantations was attributed to the richness and abundance of native understory plants (Robson et al. 2009). In the study of Lazzaro et al. (2017), they reported that the abundance and richness of microarthropods, and the richness and diversity of plant communities were significantly lower in the RP stands than these in the native oak stands, which also implies the association between understory plants and invertebrates. In this study, the RP and PC stands were characterized by greater abundance and richness of understory plants, and greater macroinvertebrate abundance in comparison to the AS stands. In combination with previous studies, the results indicate that there are positive associations between understory plants and soil macroinvertebrates. One possible reason is that the greater richness and abundance of understory plants can provide a greater array of potential habitats (Robson et al. 2009), diverse leaf-litter (Parris and Lindenmayer 2004), and alternative food resources (Salamon et al. 2011) for macroinvertebrates.
Conclusions
Our study showed that plantations of non-native Robinia pseudoacacia and Populus ×canadensis positively affected the diversity of understory plants and soil macroinvertebrates in comparison with native Armeniaca sibirica plantations. We also found that the two non-native tree plantations had inconsistent effects on specific functional groups. The results provide strong evidence that non-native plantations can affect biodiversity at different trophic levels and highlight that these levels should be considered when assessing the effects of non-native plantations on ecosystems. Given the fact that the diversity of understory plants and soil macroinvertebrates may vary among different seasons, further studies should pay attention to the seasonal variations of plantation type effects on biodiversity in the Loess Plateau.
References
Allan E, Manning P, Alt F, Binkenstein J, Blaser S, Blüthgen N, Böhm S, Grassein F, Hölzel N, Klaus VH, Kleinebecker T, Morris EK, Oelmann Y, Prati D, Renner SC, Rillig MC, Schaefer M, Schloter M, Schmitt B, Schöning I, Schrumpf M, Solly E, Sorkau E, Steckel J, Steffen-Dewenter I, Stempfhuber B, Tschapka M, Weiner CN, Weisser WW, Werner M, Westphal C, Wilcke W, Fischer M (2015) Land use intensification alters ecosystem multifunctionality via loss of biodiversity and changes to functional composition. Ecol Lett 18:834–843
Allen RB, Platt KH, Wiser SK (1995a) Biodiversity in New Zealand plantations. New Zealand For 39:26–29
Allen RB, Platt KH, Coker REJ (1995b) Understorey species composition patterns in a Pinus radiata plantation on the central North Island volcanic plateau, New Zealand. NZ J Forestry Sci 25:301–317
Bardgett RD, van der Putten WH (2014) Belowground biodiversity and ecosystem functioning. Nature 515:505–511
Bates D, Maechler M, Bolker B, Walker S (2015) lme4: linear mixed-effects models using Eigen and S4. R Package Version 1:1–9
Bertheau C, Aurélien S, Rossi JP Bankhead-dronnet S, Pineau X, Roux-morabito G, Lieutier, F (2009) Colonisation of native and exotic conifers by indigenous bark beetles (Coleoptera: Scolytinae) in France. Forest Ecol Manag 258:1619–1628
Bonham KJ, Mesibov R, Bashford R (2002) Diversity and abundance of ground-dwelling invertebrates in plantation vs. native forests in Tasmania, Australia. Forest Ecol Manag 158:237–247
Boothroyd-Roberts K, Gagnon D, Truax B (2013) Can hybrid poplar plantations accelerate the restoration of forest understory attributes on abandoned fields? Forest Ecol Manag 287:77–89
Bremer LL, Farley KA (2010) Does plantation forestry restore biodiversity or create green deserts? A synthesis of the effects of land-use transitions on plant species richness. Biodivers Conserv 19:3893–3915
Brockerhoff EG, Jactel H, Parrotta JA, Quine CP, Sayer J (2008) Plantation forests and biodiversity: oxymoron or opportunity? Biodivers Conserv 17:925–951
Chen Y, Cao Y (2014) Response of tree regeneration and understory plant species diversity to stand density in mature Pinus tabulaeformis plantations in the hilly area of the loess plateau, China. Ecol Eng 73:238–245
Chen Y, Wang K, Lin Y, Shi W, Song Y, He X (2015) Balancing green and grain trade. Nat Geosci 8:739–741
Cole L, Buckland SM, Bardgett RD (2008) Influence of disturbance and nitrogen addition on plant and soil animal diversity in grassland. Soil Biol Biochem 40:505–514
Dassou AG, Tixier P (2016) Response of pest control by generalist predators to local-scale plant diversity: a meta-analysis. Ecol Evol 6:1143–1153
Degomez T, Wagner MR (2001) Arthropod diversity of exotic vs. native Robinia species in northern Arizona. Agr Forest Entomol 3:19–27
Deng L, Shangguan ZP, Li R (2012) Effects of the grain-for-green program on soil erosion in China. Int J Sediment Res 27:120–127
Dong WH, Wang ZT, Li FR, Liu JL (2014) Characteristics of soil macrofauna community at an age sequence of cultivation in process of oasification in the middle Heihe river basin. Sci Geogr Sin 34:1254–1261
Fu BJ, Chen LD, Ma KM, Zhou H, Wang J (2000) The relationships between land use and soil conditions in the hilly area of the loess plateau in northern Shaanxi, China. CATENA 39:69–78
Gallé R, Erdélyi N, Szpisjak N, Tölgyesi C, Maák I (2015) The effect of the invasive Asclepias syriaca on the ground-dwelling arthropod fauna. Biologia 70:104–112
Gerber E, Krebs C, Murrell C, Moretti M, Rocklin R, Schaffner U (2008) Exotic invasive knotweeds (Fallopia spp.) negatively affect native plant and invertebrate assemblages in European riparian habitats. Biol Conserv 141:646–654
Gongalsky KB, Persson T, Pokarzhevskii AD (2008) Effects of soil temperature and moisture on the feeding activity of soil animals as determined by the bait-lamina test. Appl Soil Ecol 39:84–90
Gratton C, Denno RF (2005) Restoration of arthropod assemblages in a Spartina salt marsh following removal of the invasive plant Phragmites australis. Restor Ecol 13:358–372
Hartley M (2002) Rationale and methods for conserving biodiversity in plantation forests. For Ecol Manag 155:81–95
Horák J, Brestovanská T, Mladenović S, Kout J, Bogusch P, Halda JP, Zasadil P (2019) Green desert?: biodiversity patterns in forest plantations. Forest Ecol Manag 433:343–348
Hothorn T, Bretz F, Westfall P, Hothorn T, Bretz F, Westfall P (2008) Simultaneous inference in general parametric models. Biom J 50:346–363
Huang C, Zhou Z, Peng C, Teng M, Wang P (2019) How is biodiversity changing in response to ecological restoration in terrestrial ecosystems? A meta-analysis in China. Sci Total Environ 650:1–9
Huerta E, Wal HVD (2012) Soil macroinvertebrates’ abundance and diversity in home gardens in Tabasco, Mexico, vary with soil texture, organic matter and vegetation cover. Eur J Soil Biol 50:68–75
Hulme PE, Pyšek P, Jarošík V, Pergl J, Schaffner U, Vila M (2013) Bias and error in understanding plant invasion impacts. Trends Ecol Evol 28:212–218
Irwin S, Pedley SM, Coote L, Dietzsch AC, Wilson MW, Oxbrough A, Sweeney O, Moor KM, Martin R, Kelly DL, Mitchell FJ G, Kelly TC, O’Halloran J (2014) The value of plantation forests for plant, invertebrate and bird diversity and the potential for cross-taxon surrogacy. Biodivers Conserv 23:697–714
Ito S, Nakayama R, Buckley GP (2004) Effects of previous land-use on plant species diversity in semi-natural and plantation forests in a warm-temperate region in southeastern Kyushu, Japan. Forest Ecol Manag 196:213–225
Jackson ST, Hobbs RJ (2009) Ecological restoration in the light of ecological history. Science 325:567–569
Jiang Y, Kang MY, Gao QZ He LH, Xiong M, Jia ZB, Jin ZP (2003) Impact of land use on plant biodiversity and measures for biodiversity conservation in the loess plateau in China—a case study in a hilly-gully region of the northern loess plateau. Biodivers Conserv 12:2121–2133
Kasel S, Bell TL, Enright NJ, Meers TL (2015) Restoration potential of native forests after removal of conifer plantation: a perspective from Australia. Forest Ecol Manag 338:148–162
Kayler Z, Gessler A, Buchmann N (2010) What is the speed of link between aboveground and belowground processes? New Phytol 187:885–888
Kivlin SN, Hawkes CV (2011) Differentiating between effects of invasion and diversity: impacts of aboveground plant communities on belowground fungal communities. New Phytol 189:526–535
Kou M, Garcia-Fayos P, Hu S, Jiao JY (2016) The effect of Robinia pseudoacacia, afforestation on soil and vegetation properties in the loess plateau (China): a chronosequence approach. Forest Ecol Manag 375:146–158
Lazzaro L, Mazza G, D'Errico G, Fabiani A, Giuliani C, Inghilesi AF, Lagomarsino A, Landi S, Lastrucci L ,Pastorelli R, Roversi PF, Torrini G, Tricarico E, Foggi B (2017) How ecosystems change following invasion by Robinia pseudoacacia: insights from soil chemical properties and soil microbial, nematode, microarthropod and plant communities. Sci Total Environ 622–623:1509–1518
Litt AR, Cord EE, Fulbright TE, Schuster GL (2014) Effects of invasive plants on arthropods. Conserv Biol 28:1532–1549
Liu RT, Zhao HL, Zhao XY (2013) Seasonal changes of soil faunal diversity and its relation to temperature and moisture in semi-arid grassland, North China. J Arid Land Resour Environ 27:97–101
Liu RT, Zhu F, Steinberger Y (2015) Effectiveness of afforested shrub plantation on ground-active arthropod communities and trophic structure in desertified regions. CATENA 125:1–9
Meers TL, Kasel S, Bell TL, Enright NJ (2010) Conversion of native forest to exotic Pinus radiata plantation: response of understory plant composition using a plant functional trait approach. For Ecol Manag 259:399–409
Orozco-Aceves M, Standish RJ, Tibbett M (2015) Long-term conditioning of soil by plantation eucalypts and pines does not affect growth of the native jarrah tree. Forest Ecol Manag 338:92–99
Oxbrough A, Irwin S, Kelly TC, O’Halloran J (2010) Ground-dwelling invertebrates in reforested conifer plantations. Forest Ecol Manag 259:2111–2121
Oxbrough A, García-Tejero S, Spence J, O’Halloran J (2016) Can mixed stands of native and non-native tree species enhance diversity of epigaeic arthropods in plantation forests? Forest Ecol Manag 367:21–29
Paritsis J, Aizen MA (2008) Effects of exotic conifer plantations on the biodiversity of understory plants, epigeal beetles and birds in Nothofagus dombeyi forests. Forest Ecol Manag 255:1575–1583
Parris KM, Lindenmayer DB (2004) Evidence that creation of a Pinus radiata plantation in South-Eastern Australia has reduced habitat for frogs. Acta Oecol 25:93–101
Peloquin RL, Hiebert RD (1999) The effects of black locust (Robinia pseudoacacia L.) on species diversity and composition of black oak savanna/woodland communities. Nat Areas J 19:121–131
Perner J, Malt S (2003) Assessment of changing agricultural land use: response of vegetation, ground-dwelling spiders and beetles to the conversion of arable land into grassland. Agric Ecosyst Environ 98:169–181
Pimentel D, Kounang N (1998) Ecology of soil erosion in ecosystems. Ecosystems 1:416–426
Qiu LP, Zhang XC, Cheng JM, Yin XQ (2010) Effects of black locust (Robinia pseudoacacia) on soil properties in the loessial gully region of the loess plateau, China. Plant Soil 332:207–217
Quine CP, Humphrey JW (2010) Plantations of exotic tree species in Britain: irrelevant for biodiversity or novel habitat for native species? Biodivers Conserv 19:1503–1512
Roberge JM, Stenbacka F (2014) Assemblages of epigaeic beetles and understory vegetation differ between stands of an introduced pine and its native congener in boreal forest. Forest Ecol Manag 318:239–249
Robinson SI, McLaughlin ÓB, Marteinsdóttir B, OʾGorman EJ (2018) Soil temperature effects on the structure and diversity of plant and invertebrate communities in a natural warming experiment. J Anim Ecol 87: 634–646
Robson TC, Baker AC, Murray BR (2009) Differences in leaf-litter invertebrate assemblages between radiata pine plantations and neighbouring native eucalypt woodland. Austral Ecol 34:368–376
Salamon JA, Wissuwa J, Jagos S, Koblmüller M, Ozinger O, Winkler C, Frank T (2011) Plant species effects on soil macrofauna density in grassy arable fallows of different age. Eur J Soil Biol 47:129–137
Sax DF (2002) Equal diversity in disparate species assemblages: a comparison of native and exotic woodlands in California. Glob Ecol Biogeogr 11:49–57
Sitzia T, Campagnaro T, Dainese M, Cierjacks A (2012) Plant species diversity in alien black locust stands: a paired comparison with native stands across a North-Mediterranean range expansion. For Ecol Manag 285:85–91
Sylvain ZA, Buddle CM (2010) Effects of forest stand type on oribatid mite (Acari: Oribatida) assemblages in a southwestern Quebec forest. Pedobiologia 53:321–325
ter Braak CJF, Šmilauer P (2012) Canoco 5 reference manual and User's guide: software for ordination. Microcomputer Power, Ithaca
Thijs KW, Aerts R, Pieter VDM, Musila W, Gulinck H, Muys B (2014) Contrasting cloud forest restoration potential between plantations of different exotic tree species. Restor Ecol 22:472–479
Wagg C, Bender SF, Widmer F, Mg VDH (2014) Soil biodiversity and soil community composition determine ecosystem multifunctionality. Proc Natl Acad Sci U S A 111:5266–5270
Wang YF, Fu BJ, Lü YH, Chen LD (2011) Effects of vegetation restoration on soil organic carbon sequestration at multiple scales in semi-arid loess plateau, China. CATENA 85:58–66
Wardle DA, Bardgett RD, Klironomos JN, Setala H, van der Putten WH, Wall DH (2004) Ecological linkages between aboveground and belowground biota. Science 304:629–633
Willis JC, Bohan DA, Powers SJ, Choi YH, Gussin E (2008) The importance of temperature and moisture to the egg-laying behaviour of a pest slug, Deroceras reticulatum. Ann Appl Biol 153:105–115
Wu P, Zhang H, Wang Y (2015) The response of soil macroinvertebrates to alpine meadow degradation in the Qinghai–Tibetan plateau, China. Appl Soil Ecol 90:60–67
Xu M, Zhang J, Liu GB, Yamanaka N (2014) Soil properties in natural grassland, Caragana korshinskii, planted shrubland, and Robinia pseudoacacia, planted forest in gullies on the hilly loess plateau, China. CATENA 119:116–124
Yang JW, Liang ZS, Han RL (2006) Water use efficiency characteristics of four tree species under different soil water conditions in the loess plateau. Acta Ecol Sin 26:558–565
Yang Y, Jia XX, Wendrothc O, Liu BY, Shi YZ, Huang TT, Bai X (2019) Noise-assisted multivariate empirical mode decomposition of saturated hydraulic conductivity along a south-north transect across the loess plateau of China. Soil Sci Soc Am J 83:311–323
Yin WY (2001) Pictorial keys to soil faunas of China. Science Press, Beijing
Zheng LY, Gui H (2004) Insect classification. Nanjing Normal University Press, Nanjing
Zhou HJ, Van Rompaey A, Wang JA (2009) Detecting the impact of the “grain for green” program on the mean annual vegetation cover in the Shaanxi province, China using SPOT-VGT NDVI data. Land Use Policy 26:954–960
Zobel BG, Van Wyk G, Stahl P (1987) Growing exotic forests. John Wiley and Sons, New York
Acknowledgements
This research was supported by the National Key Research and Development Program of China (grant No 2016YFC0501602), the National Natural Science Foundation of China (grant No 41671271), and the China Postdoctoral Science Foundation (2018 M641491). We thank reviewers for helpful comments on the manuscript.
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Zhu, Y., Wang, Y. & Chen, L. Effects of non-native tree plantations on the diversity of understory plants and soil macroinvertebrates in the Loess Plateau of China. Plant Soil 446, 357–368 (2020). https://doi.org/10.1007/s11104-019-04336-y
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DOI: https://doi.org/10.1007/s11104-019-04336-y