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Published online December 12, 2024
https://doi.org/10.5141/jee.24.076

Journal of Ecology and Environment (2024) 48:46

Roles of Chromolaena odorata, macrofauna, and forest edge on the decomposition rate of tree leaf litter in two types of seasonally dry tropical forest

Kasidit Rison1 , Marut Fuangarworn2,3 and Chatchawan Chaisuekul2,3*

1Ph.D. Program in Zoology, Department of Biology, Faculty of Science, Chulalongkorn University, Bangkok 10330, Thailand
2Integrative Insect Ecology Research Unit, Department of Biology, Faculty of Science, Chulalongkorn University, Bangkok 10330, Thailand
3Department of Biology, Faculty of Science, Chulalongkorn University, Bangkok 10330, Thailand

Correspondence to:Chatchawan Chaisuekul
E-mail Chatchawan.c@chula.ac.th

Received: August 19, 2024; Revised: October 24, 2024; Accepted: November 11, 2024

This article is licensed under a Creative Commons Attribution (CC BY) 4.0 International License, which permits use, sharing, adaptation, distribution and reproduction in any medium or format, as long as you give appropriate credit to the original author(s) and the source, provide a link to the Creative Commons license, and indicate if changes were made. The images or other third party material in this article are included in the article’s Creative Commons license, unless indicated otherwise in a credit line to the material. If material is not included in the article’s Creative Commons license and your intended use is not permitted by statutory regulation or exceeds the permitted use, you will need to obtain permission directly from the copyright holder. To view a copy of this license, visit http://creativecommons.org/licenses/by/4.0/ The publisher of this article is The Ecological Society of Korea in collaboration with The Korean Society of Limnology

Abstract

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Background: The forest edge of seasonally dry tropical forests (SDTF) is commonly invaded by Chromolaena odorata, which could affect the leaf-litter decomposition (LLD) rate through the litter fauna and the litter chemistry. We tested the effects of positions (edge vs. interior), C. odorata inclusion, and macrofauna inclusion using litterbags containing the two most dominant tree species from each forest type of SDTF (a dipterocarp deciduous forest and a mixed deciduous forest [MDF]), in northern Thailand.
Results: The results showed that leaf litter decayed at the same rate between the forest edge and interior. A difference in the decomposition rate between the edge and interior was only found in the MDF and only when the composition of the litter contained a high nitrogen (C. odorata) and phosphorous content (Pterocarpus macrocarpus – a native tree of MDF). Exclusion of macrofauna resulted in slower decomposition rates, but this effect was not significant when combined with the edge condition.
Conclusions: Our findings indicated that the forest edge condition has no directly significant effect on the LLD rate but is mediated through changes in the litter chemistry by inclusion of C. odorata litter.

Keywords: forest edge, invasive herbaceous species, leaf-litter decomposition, seasonally dry tropical forest

Introduction

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Seasonally dry tropical forests (SDTF) are a widespread forest ecosystem in southeastern Asia (Dexter et al. 2015) that have been subject to significant degradation and fragmentation due to anthropogenic activities (Delang 2002; Wanthongchai et al. 2008). These disturbances have led to the emergence of forest edges, which create microclimate variations between the interior and exterior of forest borders (Ewers and Didham 2006; Laurance et al. 2011). The forest edge causes a significant change in the forest biodiversity, such as increased levels of herbaceous plants (Szigeti et al. 2022) and soil fauna (De Smedt et al. 2016; De Smedt et al. 2019; Lacasella et al. 2015), which can lead to several adverse impacts on the forest ecosystem functions (Ewers et al. 2011; Sodhi et al. 2010).

Leaf-litter decomposition (LLD) is a critical process that contributes to carbon (C) and nutrient cycling and so ecosystem productivity in tropical forest ecosystems (Hättenschwiler et al. 2005; Zhang and Zak 1995). The decomposition of leaf litter is a complex process that is mediated by both biotic and abiotic factors (Cornwell et al. 2008; Krishna and Mohan 2017). In tropical forests that share a similar climate, variations in the LLD rate are primarily due to the chemical composition of the litter species and the activity of soil organisms (García-Palacios et al. 2013; González and Seastedt 2001; Meyer et al. 2020; Njoroge et al. 2023). It is known that these two factors are possibly altered along the edge-interior gradient in tropical forests (De Smedt et al. 2016; Salles et al. 2018).

Litter faunae, mainly arthropods, on the forest floor are categorized depending on their size into macro- and meso-fauna, and these could have different effects on the ecosystem function. In tropical forests, macrofaunae are the most important group of detritivores that control the LLD rates (González and Seastedt 2001). The activity and density of these groups are known to be sensitive to habitat changes (Wang et al. 2023), such as at forest edges, where soil moistures are generally low (Birkhofer et al. 2015; Didham 1998; Ferguson 2004). When the distance from the forest edge in a dry forest increases, the canopy cover and soil humidity decrease (Dambros et al. 2013). These drier conditions may restrict the activity of litter macro-arthropods, which could retard the rate of LLD caused by these organisms (Didham 1998; Paudel et al. 2015).

Forest edges may also impact the litter quality by promoting the recruitment of numerous invasive herbaceous species, such as Chromolaena odorata, where such invasive species have been frequently reported in southern and southeastern Asia, especially in dry forests (Sharma et al. 2022; Wei et al. 2017). The presence of leaf litter from C. odorata could affect the chemical composition of the litter near the forest edges (Banful et al. 2008; Mboukou-Kimbatsa et al. 2007) because its leaves show a high nitrogen (N) content while the leaves of native tree species in SDTF generally have a low N content that can take more than a year to decompose (Hanpattanakit and Chidthaisong 2012). Mixing leaf litter from species that differ in their litter quality and structure changes the physio-chemical properties of the decomposition environment via nutrient transfer from high-quality (labile) litter to low-quality (recalcitrant) litter, resulting in an increased LLD rate (Bonanomi et al. 2014; Yang et al. 2022). Moreover, the mixed leaf litter can create more heterogeneous microenvironments due to leaf toughness and debris size, which can affect soil fauna activity and subsequently impact decomposition rates (Njoroge et al. 2022; Wang et al. 2024). As in SDTFs, which contain the majority of tree species and a few herbaceous species at the understory level (Bunyavejchewin 1983; Bunyavejchewin et al. 2011; Myo et al. 2016), forest edges heavily invaded by C. odorata have altered soil properties, such as soil moisture and organic matter (OM) content (Koné et al. 2012; Norgrove et al. 2000; Wei et al. 2017).

In Thailand, SDTFs are mainly dominated by deciduous tree species (Bunyavejchewin 1983; Bunyavejchewin et al. 2011). Mixed deciduous forests (MDF) and dipterocarp deciduous forest (DDF) are two commonly found types in northern Thailand that exist at similar elevations and experience regular disturbances from forest fires and human activities (Wanthongchai et al. 2008). The two forest types show a different species dominance (Bunyavejchewin 1983; Bunyavejchewin et al. 2011). Moreover, previous research has suggested that the soil environment of these two forest types is distinct, with MDFs showing a higher moisture and OM content in the soil while DDFs show a more sandy soil with less moisture, which could impact both the distribution and composition of the soil fauna and herbaceous species, leading to a different LLD rate between the two forest types in the same climatic region (Asanok et al. 2020; Myo et al. 2016).

Despite these potential factors that suggest that the LLD rates in fragmented SDTFs could be affected by the forest edges through the distribution of soil fauna and composition litter chemistry, there has been insufficient empirical studies to support or refute this. Therefore, we studied how the forest edge affects the LLD rate in a STDF by examining the relationships between the litter fauna and plant leaf chemistry in a MDF and DDF in northern Thailand. The experiment was conducted over 18 months as a litter bag experiment, which considered the contribution of herb litter and access to the litter fauna between the forest edge and interior. We hypothesized that the forest edge changes the composition of the leaf litter and litter fauna and these in turn increase the LLD rate compared to in the forest interior.

Materials and Methods

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Study site

The study was carried out in a 350-ha SDTF in Nan province, northern Thailand (18.55 N, 100.79 E, 200 m above mean sea level). The climate is highly seasonal, with the rainy season lasting six months from May to October and the dry season lasting from December to mid-April. The precipitation in the area was 2,314 mm on an annual average basis, with a minimum–maximum temperature of 21.3°C–35.2°C. The dry season has an average temperature of 35.3°C and rainfall of 312.5 mm, whereas the rainy season has an average temperature of 29.4°C and rainfall of 1,935 mm. The area’s forests are classified into two types of SDTFs – MDF and DDF – according to their zone of distribution. The DDF is located in the east part of the study site and covers about 200 ha, whereas the MDF is located in the west part of the site (Fig. 1) (Dumrongrojwatthana 2004). Pterocarpus macrocarpus, Aporosa villosa, and Xylia xylocarpa are the most prominent tree species in the MDF, whereas Shorea obtusa and Dipterocarpus tuberculatus are the dominant species in the DDF (Dumrongrojwatthana 2004). Around the fragment, various anthropocentric lands surround the forest remnants, including residential development, agriculture, abandoned fields, and transportation infrastructure.

Figure 1. Maps of the study area of a fragmented seasonally dry tropical forest in Nan province, northern Thailand (A, B). Shown are the two forest types and location of the study transects (C), landscape of forest edge (D), and experimental plot (E). DDF stands for dipterocarp deciduous forest and MDF stands for mixed deciduous forest.

Litter bag experiment

The leaf litter from the two most dominant tree species were selected from each type of forest (P. macrocarpus and A. villosa for MDF; and S. obtusa and D. tuberculatus for DDF) along with the herbaceous C. odorata from the forest edge. During the leaf-falling season, around February to March 2020, freshly senesced leaf material from these five species were collected within the forest remnant. The collected litter was dried at 80°C for 48 hours and then stored at 4°C before mixing at the given proportion and filling in litter bags. All samples of the five species were analyzed for their chemical components, including C, N, potassium (K), and phosphorous (P). The C and N contents were analyzed using the Dumas method with a CN Analyzer (CN CORDER MT-700; Yanaco, Kyoto, Japan), while the P and K contents were evaluated using the wet ashing method with a 5:1:2 molar ratio of HNO3: H2SO4: HClO4. The analysis of P (vanadomolybdate yellow color) was conducted at a wavelength of 440 nm in a spectrophotometer, while the analysis of K was performed using atomic absorption spectrophotometry.

The dried leaf-litter were enclosed in mesh bags (20 × 20 cm2, 30 g of litter weight) and placed on the ground, where they were collected at 5 periodic intervals (3, 6, 9, 12, and 18 months) to determine the remaining mass. Two mesh sizes of nylon-litter bags were chosen: 2 mm (micro- and meso-fauna pass through) and 5 mm (micro-, meso-, and macro-fauna pass through) (Dossa et al. 2016; Paudel et al. 2015). The litter quality (variations in chemical components of each species) along each transect was ascertained using the two dominant tree species for each forest type and one herbaceous species. Therefore, the litter bags contained tree litter (1:1 weight mixture of two tree species) and mixed herbaceous litter (1:1:1 weight mixture of leaves from the two dominant tree species and C. odorata). Leaves of D. tuberculatus were trimmed into 20 × 20 m2 to fit into the litter bags, while the other leaf species remained untrimmed.

Five transects from each forest type were selected in the study site and were located 300 m away from each other. Within each transect, two 10 × 10 m2 plots were set up in the field, one was located near the forest edge (0–5 m from the physical forest edge), and another were located in the forest interior (100–120 m from the physical forest edge). The environmental factors (soil moisture, canopy, and C. odorata cover, leaf litter, and the soil type, N, and C content) were ascertained in the plots within August of the year 2020 and 2021. The average soil moisture, C, and N content was determined at the four corners and center using a 100 cm3 volume of the collected topsoil layer. The soil OM (Walkley and Black 1934) and N (Bremner 1960) were measured by Department of Soil Science, Faculty of Agriculture, Kasetsart University, Thailand. The fresh to dry weight ratio was used to determine the soil moisture content. Densitometers were used to determine the average canopy cover per plot at the same location as the previous measurement. Five quadrats (20 × 20 cm2) per plot were used to collect the leaf litter on the forest floor, then dried and weighed. The C. odorata cover was estimated as an average from three subplots of 1 × 1 m2.

Litter bags were placed on the soil surface at center of each plot (10 × 10 m2). We designed an evenly spaced four rows of five column sampling-grids and placed one litterbag for each mesh size and each plant species at each sampling point along the grid. A total of 400 litter bags (two types of plant species combination × two mesh sizes × two locations of plot × two forest types × five replicated plots × five sampling months) were placed in the forest. Litter bags were spread as flat as possible to maintain contact with the soil during the entire decomposition period in the incubation plots. To mimic natural decomposition conditions, litter bags were covered with a natural litter layer (Paudel et al. 2015). To avoid the litter bags being carried away by wind, a large mesh net was (12.7 cm mesh size) was laid on top of the litter bags after being placed on the soil surface. During the experimental period, a total of 31 of the 2-mm litter bags were removed from the analysis due to traces of termite destruction.

A set of eight litter bags (two plant species treatments × two mesh sizes × two locations) was removed from each transect for each of five sampling months (3, 6, 9, 12, and 18 months) and brought to the lab in a sealed plastic bag for further analysis. The outside of the litter bags was thoroughly inspected, and any foreign debris or plant matter was removed using forceps. The macrofauna found visually were collected by hand, and then the litter was extracted for fauna (macro- and meso-fauna) using Tullgren funnels. A 70% (v/v) ethanol solution was used to collect the soil fauna samples. The samples were sorted into broad taxonomic categories and counted under a stereoscopic light microscope. The relative density of the soil fauna was determined by comparing the number of groups and individuals collected from particular litterbags to the total dry weight of the litter in those bags. Following removal of the soil fauna, the litter was thoroughly rinsed with distilled water to remove any remains of soil or organic material. After being oven-dried at 60°C for 72 hours, the remaining mass of litter in all the collected litter bags was measured.

Data analysis

Differences of environmental factors between the forest edge and forest interior were examined using the paired samples the related samples Wilcoxon signed rank test due to the data were asymmetrically distributed.

Initial litter weight (IL) and percentages of remaining litter weight (%RL) were calculated using a comparison of the weight of litter inside each bag prior to placement in the field at the onset of experiment. The RL was the weight of litter in each bag after each extraction was measured. The %RL, determined as shown in equation 1, was then used to determine the mean litter decomposition rate (k) for each forest site using the model (Olson 1963) as shown in equation 2.

% Remaining litter(g)=Remaining litterInitial Litter×100
Remaining litter=Initial litter×e−kt

where t is the time in years and k is the decomposition rate.

This exponential model was linearized using the natural logarithm of the %RL in order to calculate the linearized decomposition rate, as shown in equation 3,

k=−ln(RL/IL)/t

All statistical analyses were separated between forest types and were performed using the R program (R Core Team 2022). Using the “nls” function of the R stats packages, the k value for LLD was calculated using sets of data representing replicates and incubation times for each treatment. We assessed normality using the “shapiro.test” in stats packages and homoscedasticity using the “bp.test” function (Zeileis and Hothorn 2002). Ordinary least square tests were conducted to determine if the initial leaf-litter characteristics could predict the k values for these treatments. We accounted for variations in k values for litter decomposition using multiple linear regression models comprised of the compositions of litter species, forest edge, mesh size of litter bags, and their interactions. The analysis was separated between forest types.

Results

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Leaf litter chemistry and forest environmental factors

The C. odorata litter contained a significantly higher N content than the other four litter types (S. robusta, D. tuberculatus, P. macrocarpus, and A. villosa) but a similar amount of C as P. macrocarpus and D. tuberculatus litters (Table 1). The A. villosa and D. tuberculatus liters had more organic C than the other litters, while S. robusta had the lowest amount. The C. odorata and P. macrocarpus litter had a lower C:N ratio than the other litters, whereas S. robusta, D. tuberculatus, and A. villosa litter had a higher C:N ratio (Table 1).

Table 1 . Initial concentration of mixed litter in two forest types.

Litter speciesOM (%)Organic C (%)N (%)P (%)K (%)C:N ratio
Shorea obtusa55.49 (0.59)32.19 (0.12)0.37 (0.49)0.02 (0.25)0.3 (0.11)60.74 (0.22)
Dipterocarpus tuberculatus58.68 (0.05)34.04 (0.15)0.31 (0.37)0.03 (0.19)0.27 (0.94)77.36 (0.32)
Pterocarpus macrocarpus57.26 (0.33)33.21 (0.34)0.37 (0.13)0.24 (0.08)1.87 (0.09)63.87 (0.29)
Aporosa villosa62.85 (0.13)36.46 (1.15)0.35 (0.07)0.03 (0.39)0.17 (0.07)72.92 (0.04)
Chromolaena odorata56.6 (0.24)34.47 (0.03)1.17 (0.04)0.08 (0.69)0.13 (0.34)20.89 (0.04)

The leaf-litter species include native and invasive ones. Values in parenthesis are the standard errors.

OM: organic matter; C: carbon; N: nitrogen; P: phosphorous; K: potassium.

The comparison of forest environmental factors between the forest edge and forest interior indicated that the leaf litter at forest floor and C. odorata cover was more pronounced at the forest edge. Conversely, canopy cover, OM, total N and soil water content exhibited no significant differences between the forest edge and the forest interior. This trend was observed consistently across both forest types (Table 2).

Table 2 . Environmental factors between the edge and interior of two forest types.

Environmental propertiesDDFMDF
Forest edge (0–5 m)Forest interior (~120 m)Forest edge (0–5 m)Forest interior (~120 m)
Canopy cover65.43 (2.56)80.32 (4.23)67.44 (4.32)86.12 (3.94)
Soil water content26.70 (5.68)25.02 (0.96)35.55 (2.18)38.03 (2.56)
Organic matter (g/kg)10.1 (1.23)9.7 (1.90)14.2 (1.11)13.0 (1.75)
Total N (%)0.48 (0.09)0.39 (0.05)0.73 (0.05)0.65 (0.03)
Floor leaf litter (dried weight, g/m2)125.32 (5.34)a105.11 (3.78)a97.45 (6.41)a85.64 (5.44)a
Chromolaena odorata cover (%)23.3 (2.12)a6.5 (1.86)a26.3 (3.10)a3.2 (0.73)a

Values in parenthesis are the standard errors.

DDF: dipterocarp deciduous forest; MDF: mixed deciduous forest; N: nitrogen.

aSignificant difference at p < 0.05.

Litter faunae between forest edges and interior

A total of 24 distinct soil fauna groups were recorded in the litter bags. Acari comprised 47.2%–55.2% of the total number of litter fauna individuals across the forest types and forest edge location. Collembola, Hymenoptera (ants), and Isoptera were, respectively, the second, third, and fourth most abundant components of the litter fauna in each of the two forest types. Compared to the 2-mm mesh bags, the 5-mm litter bags decreased the individual abundance of total fauna (individuals per plot) by 24.31% and 20.46%, respectively, in the DDF and MDF as well as the taxonomic diversity.

Leaf litter mass loss and decomposition rate (k)

The 18-month litter mass loss ranged from 33 to 62% of the total mass loss. The logarithmic regressions of mass loss for each treatment (mesh size, contribution of C. odorata, and forest edge) and forest type all showed good fits (all p < 0.001), indicating that the use of a single exponential model for calculation and comparison was reasonable. The mesh size, forest edge, and contribution of C. odorata and their interactions all significantly affected the litter k (Figs. 2 and 3). In the two forest types, the application of 5-mm mesh bags resulted in a higher mass loss at the end of the incubation period (month 18) than in the 2-mm mesh bags (16.6% in DDF and 17.2% in MDF). In addition, there were significant effects of mixing in C. odorata in only the MDF (DDF: t = 0.656, p = 0.516; MDF: t = 2.104, p = 0.043). The location of litter bags installed at the forest edges had no significant difference in the k value (DDF: t = –1.219, p = 0.231; MDF: t = –0.180, p = 0.858). However, the treatment interaction between the forest edge and litter mixing was found to be significantly different only in the MDF, but not in the DDF (DDF: t = 0.425, p = 0.673; MDF: t = 2.001, p = 0.050) (Table 3).

Table 3 . Summary of multiple regression analysis, showing the effect of forest type, location, mesh size, and presence of Chromolaena odorata on the linearized k-rate for the two different forest types.

TreatmentsDDFMDF
tp-valuetp-value
Forest edge–1.2190.231–0.1800.858
Mesh size (5-mm)7.073< 0.017.447< 0.01
Mixtures with C. odorata0.6560.5162.1040.043
Mesh size (5-mm) × forest edge–0.1090.913–1.5910.121
Mixtures with C. odorata × forest plot0.4250.6732.0010.050
Mesh size (5-mm) × mixtures with C. odorata0.0190.9840.2310.818
Mesh size (5-mm) × mixtures with C. odorata × forest edge–0.1190.905–0.1900.850

DDF: dipterocarp deciduous forest; MDF: mixed deciduous forest.

Figure 2. Litter mass loss (% initial dry mass) of different species in litter bags with different mesh sizes and litter mixture across sampling sites. (A) Forest edge of DDF. (B) Forest interior of DDF. (C) Forest edge of MDF. (D) Forest interior of MDF. Values are the means ± standard error, and a logarithmic fitting for each treatment is given. DDF stands for dipterocarp deciduous forest and MDF stands for mixed deciduous forest.

Figure 3. Effects of tree species, litter bag mesh size, and their interactions on the leaf litter decomposition rates (k, year−1) in DDF (A) and MDF (B). Values are means ± standard deviation, and different letters indicate significant (p < 0.05) differences between treatments. DDF stands for dipterocarp deciduous forest and MDF stands for mixed deciduous forest.

Discussion

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Effect of forest edge on the community of fauna

Overall, the litter fauna density and assemblage were different across the two forest types. This indicated the important effect of tree species on the soil environment and microclimate in a similar climatic region (Li et al. 2022; Seidelmann et al. 2016; Tedersoo et al. 2016), which in turn affects the density of some fauna groups. However, within each forest type, no differences between the two plot locations (forest edge and forest interior) were observed (except for Acari). This was because there was very little change in the abiotic variables (i.e., air temperature, evaporation rate, and moisture content) due to the forest edge interior gradient. Similarly, previous studies on the succession of tropical forests revealed that the forest’s physical environment due to forest edge had little or no effect on the detritivore community (Moreno et al. 2014; Moreno et al. 2020). Out of various types of disturbances, only high disturbance levels, such as anthropogenic clearance (Bloemers et al. 1997; Coyle et al. 2017; Paudel et al. 2015), resulted in microclimate changes that were significant enough to limit the activity of detritivores species in tropical forests (Coyle et al. 2017; Dossa et al. 2016; Paudel et al. 2015).

The presence of C. odorata has been shown to have significant impacts on soil properties and microbial activities (Koné et al. 2021). In invaded areas, soil microbial biomass carbon, carbon mineralization, enzymatic activities, and available N and phosphorus generally increased compared to non-invaded areas (Koné et al. 2021). This suggests that a lower coverage of C. odorata in forest interiors might result in reduced soil microbial activity and nutrient availability compared to forest edges with higher invasion. Moreover, the effects of C. odorata on soil properties appear to be more pronounced in MDF compared to DDF. MDF sites generally have higher soil fertility, with greater soil moisture, OM, total N, available phosphorus, and exchangeable K than DDF sites (Charoonphong and Suwanwaree 2014). This higher initial fertility in MDF may provide a more favorable environment for C. odorata to establish. The litter quality of C. odorata has a low lignin/N ratio (Mboukou-Kimbatsa et al. 2007), which can lead to faster decomposition and nutrient release (Koné et al., 2021), as soil conditions are generally more favorable for soil fauna. A study showed that the presence of C. odorata increases earthworm density and promotes soil aggregate formation (Mboukou-Kimbatsa et al. 2007). The increased faunal activity can enhance litter incorporation into the mineral soil, potentially leading to greater changes in soil organic C distribution in MDF compared to DDF (Guidi et al. 2022).

Effect of litter fauna on the leaf-litter decomposition rates

Many previous studies have shown that the macrofauna had a higher impact on the LLD rates than the mesofauna did (Hättenschwiler et al. 2005; Peguero et al. 2019; Yang and Chen 2009). Our findings here showed that the LLD rates were faster when the macrofauna could gain access to the litter bags. However, the effect is not pronounced with respect to the forest edge. As termites were the majority macrofauna taxa found in this study and were present at the same density in both the forest edge and interior, the presence of the forest edge did not shift the LLD rate in our experiment regardless of the litter species chemistry. Termites can survive in dry conditions (Ashton et al. 2019), and indeed Macrotermes thrive in this condition (Arshad 1982). Moreover, the supplementing of leaf litter with C. odorata litter may not alter the termite feeding, as several studies have shown that termites show only a slight preference for the leaf litter of different species (Matsumoto and Abe 1979).

Macro- and meso-fauna responses differ from habitat conversion, such as forest edges, where the macrofauna density has been reported to be more affected by the habitat variation than the mesofauna, but the mesofauna community are more sensitive than the macrofauna (Wang et al. 2023). Our result shows the opposite trend, where the litter mesofauna contributed to the high density at the forest edge plot because colonization of herbaceous species shifted the soil physical factors, such as the soil moisture, N, and C contents, to provide a better food source and habitat complexity in the soil and litter layer (Sabatté et al. 2021; Zheng et al. 2022). As in SDTFs, which contain the majority of tree species and a few herbaceous species at the understory level (Bunyavejchewin 1983; Bunyavejchewin et al. 2011), forest edges heavily invaded by C. odorata alter the soil properties, such as the soil moisture and OM content (Koné et al. 2012; Norgrove et al. 2000; Wei et al. 2017), resulting in a higher density of Acari at the MDF edge. In contrast, the soil in the DDF was sandier than in the MDF (Myo et al. 2016), which may explain the reduced levels of C, N, and moisture, and so making it unfavorable for soil mesofauna.

Even though Acari are frequently mentioned as dominant fauna in litterbag experiments (Xin et al. 2012) and exhibit significant variations in response to environmental factors (Ahmed et al. 2020; Elmoghazy and Shawer 2013; Manu 2011), their contribution to decomposition appears limited. Because the feeding guilds of soil mites in tropical forests span a wide range—including detritivores, fungivores, omnivores, and predators (Díaz-Aguilar and Quideau 2013)—not all mites directly contribute to litter decomposition. The study on OM breakdown found no significant treatment-related effects when Acari abundances were reduced by up to 80% using insecticides (Pamminger et al. 2022). This suggests that other trophic levels, such as microorganisms or termites, may play a more crucial role in decomposition than the mesofauna community in tropical forests (Abe 2019; Fujii et al. 2016).

Seasonal variations also play a role in litter decomposition and fauna activity in tropical forests (González and Seastedt 2001), which could significantly impact decomposition rates in this study, regardless of the effect of litter quality. Tropical arthropods exhibit very high rates of intra-annual turnover in species composition, associated with seasonal changes in environmental variables (Beng et al. 2018) such as temperature, rainfall, and humidity (Krishna and Mohan 2017). The maximum leaf litter breakdown rate coincided with the rainy season (de Souza Rezende et al. 2016).

Effect of adding C. odorata on the leaf-litter decomposition rate

The results showed that the inclusion of C. odorata into the leaf litter did not change the LLD rate of the DDF tree leaf litter (S. robusta and D. tuberculatus) but sped up that of MDF tree leaf litter (A. villosa and P. macrocarpus) by approximately 15%. Despite the stronger influence on the decomposition rate of low-quality litter seen in many previous studies (Fuqiang et al. 2010; Gartner and Cardon 2004; Steinwandter et al. 2019), the regulation of the LLD rates of the tree species was not affected by the mixing of high- and low-quality litter. Thus, adding high-quality litter from a different species may not necessarily drive the decomposition rates of the target litter (Santonja et al. 2017), as demonstrated by the opposite effects of adding C. odorata litter on the decomposition of tree litter, even though the soil contained a lower N content than in the leaf litter. The litter’s potential regulating effect on the N and P concentrations and other environmental factors may be the cause (Feller et al. 2003). Rapid decomposition and a high initial concentration of N were observed in the C. odorata litter in the MDF due to the combined effect between the N and P concentrations. Mixing both C. odorata and P. macrocarpus leaf litter resulted in a significant rise in the P concentration in the total litter chemistry, which may have had a direct and positive impact on the LLD rate. Previous research has identified consistent patterns, where a higher N input only resulted in the inhibition of the decomposition of low-quality litter (Knorr et al. 2005), while a combination of N and P enrichment caused an enhanced LLD (Ochoa-Hueso et al. 2019; Tie et al. 2023).

Nutrient transfer dynamics between N and P differ significantly during decomposition. Under laboratory conditions, C. odorata attained 100% decomposition within 8 weeks (Okeke and Omaliko 1992). N from C. odorata could be rapidly released and depleted from the litter. Phosphorus is often strongly retained and released more slowly, particularly in recalcitrant litters (Ong et al. 2017; Robbins et al. 2019). The pattern of rapid N depletion and slower P release was observed in other study. In tropical peat swamp forests, leaf litter P content was rapidly depleted, while N release was more gradual (Ong et al. 2017).

Furthermore, our findings suggest that the antagonistic effects observed in DDF species may be attributable to the high lignin content of D. tuberculatus and S. robusta litter compared to that of P. macrocarpus (Osono 2020; Sarjubala Devi and Yadava 2007; Yang et al. 2020). Protein complexes, like those produced by lignin, are difficult for most organisms to degrade (Hättenschwiler and Vitousek 2000).

Effect of forest edge on leaf-litter decomposition between forest types

The MDF was suggested to have higher decomposition rates due to more favorable conditions for decomposer organisms, such as higher soil nutrient content and greater plant diversity (Asanok et al. 2022; Ishida et al. 2023). Moreover, the leaf traits of the MDF are dominated by species with higher specific leaf area and leaf dry matter content, while the DDF is characterized by species with higher wood density and thicker leaves (Asanok et al. 2022). Higher specific leaf area is generally associated with faster decomposition rates, which could indicate more rapid decomposition. Additionally, the DDF experiences more frequent fires compared to the MDF (Khaing et al. 2019). Fire can also affect decomposition rates by altering soil properties and vegetation structure, as well as the amount of C transferred to the mineral soil (Campo and Merino 2016). This association between increased fire frequency and reduced decomposition rates could, in turn, impact the differences in C storage between the DDF and MDF (Campo and Merino 2016;, Mondal and Sukumar 2013).

Conclusions

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The forest edge influenced the LLD rate of the native tree species of a SDTF in certain conditions, specifically by adding a higher N content in the litter composition (addition of C. odorata to P. macrocarpus litter). However, this effect was only found in the leaf litter that was not mixed with the hard to decompose species, such as S. robusta and D. tuberculatus. Moreover, the macrofauna had a robust impact on the LLD rate regardless of the forest edge conditions. Our finding emphasizes the importance of litter species in shaping the LLD under a forest edge condition to gain a better understanding of the impact of forest fragmentation on ecosystem function.

Acknowledgements

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We are grateful to Chulalongkorn University’s Center of Learning Network for the Region in Nan province for providing research facilities and accommodation during the study period.

Authors’ contributions

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KR conceived the ideas, conducted field study, drafted the manuscript, designed the figures, and analyzed the data and results. MF and CC conceived the ideas and reviewed the manuscript. All authors discussed the results and commented on the manuscript.

Funding

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The Plant Genetic Conservation Project Under the Royal Initiative of Her Royal Highness Princess Maha Chakri Sirindhorn (RSPG-CU) provided financial support for this study.

References

Go to
  1. Abe T. In: Breed MD, Michener CD, Evans HE, editors. The biology of social insects. Boca Raton: CRC Press; 2019. p. 71-5.
    KoreaMed CrossRef
  2. Ahmed R, Gupta SK, Roy S, Bora D. Diversity and seasonal variation of mesostigmatid mites in three tea gardens of Assam (India) with different agro-practices. Proc Entomol Soc Wash. 2020;122(4):750-6. https://doi.org/10.4289/0013-8797.122.4.750.
    CrossRef
  3. Arshad MA. Influence of the termite Macrotermes michaelseni (Sjöst) on soil fertility and vegetation in a semi-arid savannah ecosystem. Agro-Ecosystems. 1982;8(1):47-58. https://doi.org/10.1016/0304-3746(82)90014-2.
    CrossRef
  4. Asanok L, Taweesuk R, Kamyo T. Edge tree functional traits and their association with edaphic factors in seasonally dry forests in northern Thailand. iForest. 2022;15(4):273-80. https://doi.org/10.3832/ifor3870-015.
    CrossRef
  5. Asanok L, Taweesuk R, Papakjan N. Woody species colonization along edge-interior gradients of deciduous forest remnants in the Mae Khum Mee watershed, Northern Thailand. Int J For Res. 2020;2020:1-13. https://doi.org/10.1155/2020/5867376.
    CrossRef
  6. Ashton LA, Griffiths HM, Parr CL, Evans TA, Didham RK, Hasan F, et al. Termites mitigate the effects of drought in tropical rainforest. Science. 2019;363(6423):174-7. https://doi.org/10.1126/science.aau9565.
    Pubmed CrossRef
  7. Banful B, Ofori K, Kumaga F, Hauser S, Ndango R. Decomposition and nutrient release patterns of Pueraria phaseoloides, Flemingia macrophylla and Chromolaena odorata leaf residues in tropical land use systems. Ghana Jnl Agric Sci. 2008;41(1):11-22. https://doi.org/10.4314/gjas.v41i1.46141.
    CrossRef
  8. Beng KC, Corlett RT, Tomlinson KW. Seasonal changes in the diversity and composition of the litter fauna in native forests and rubber plantations. Sci Rep. 2018;8(1):10232. https://doi.org/10.1038/s41598-018-28603-7.
    Pubmed KoreaMed CrossRef
  9. Birkhofer K, Smith HG, Weisser WW, Wolters V, Gossner MM. Land-use effects on the functional distinctness of arthropod communities. Ecography. 2015;38(9):889-900. https://doi.org/10.1111/ecog.01141.
    CrossRef
  10. Bloemers GF, Hodda M, Lambshead PJ, Lawton JH, Wanless FR. The effects of forest disturbance on diversity of tropical soil nematodes. Oecologia. 1997;111(4):575-82. https://doi.org/10.1007/s004420050274.
    Pubmed CrossRef
  11. Bonanomi G, Capodilupo M, Incerti G, Mazzoleni S. Nitrogen transfer in litter mixture enhances decomposition rate, temperature sensitivity, and C quality changes. Plant Soil. 2014;381:307-21. https://doi.org/10.1007/s11104-014-2119-4.
    CrossRef
  12. Bremner JM. Determination of nitrogen in soil by the Kjeldahl method. J Agric Sci. 1960;55(1):11-33. https://doi.org/10.1017/S0021859600021572.
    CrossRef
  13. Bunyavejchewin S. Analysis of the tropical dry deciduous forest of Thailand, I. Characteristics of the dominance-types. Nat Hist Bull Siam Soc. 1983;31(2):109-22.
  14. Bunyavejchewin S, Baker PJ, Davies SJ. In: McShea WJ, Davies SJ, Bhumpakphan N, editors. The ecology and conservation of seasonally dry forests in Asia. Washington, D.C.: Smithsonian Institution Scholarly Press; 2011. p. 9-35.
  15. Campo J, Merino A. Variations in soil carbon sequestration and their determinants along a precipitation gradient in seasonally dry tropical forest ecosystems. Glob Chang Biol. 2016;22(5):1942-56. https://doi.org/10.1111/gcb.13244.
    Pubmed CrossRef
  16. Charoonphong W, Suwanwaree P. The study of soil physical and chemical properties in Plant Genetic Protection Area of RSPG, Nampung Dam EGAT, Thailand. Adv Mater Res. 2014;911:393-6. https://doi.org/10.4028/www.scientific.net/AMR.911.393.
    CrossRef
  17. Cornwell WK, Cornelissen JH, Amatangelo K, Dorrepaal E, Eviner VT, Godoy O, et al. Plant species traits are the predominant control on litter decomposition rates within biomes worldwide. Ecol Lett. 2008;11(10):1065-71. https://doi.org/10.1111/j.1461-0248.2008.01219.x.
    Pubmed CrossRef
  18. Coyle DR, Nagendra UJ, Taylor MK, Campbell JH, Cunard CE, Joslin AH, et al. Soil fauna responses to natural disturbances, invasive species, and global climate change: current state of the science and a call to action. Soil Biol Biochem. 2017;110:116-33. https://doi.org/10.1016/j.soilbio.2017.03.008.
    CrossRef
  19. Dambros CS, da Silva VNV, Azevedo R, de Morais JW. Road-associated edge effects in Amazonia change termite community composition by modifying environmental conditions. J Nat Conserv. 2013;21(5):279-85. https://doi.org/10.1016/j.jnc.2013.02.003.
    CrossRef
  20. De Smedt P, Baeten L, Proesmans W, Van de Poel S, Van Keer J, Giffard B, et al. Strength of forest edge effects on litter-dwelling macro-arthropods across Europe is influenced by forest age and edge properties. Divers Distrib. 2019;25(6):963-74. https://doi.org/10.1111/ddi.12909.
    CrossRef
  21. De Smedt P, Wuyts K, Baeten L, De Schrijver A, Proesmans W, De Frenne P, et al. Complementary distribution patterns of arthropod detritivores (woodlice and millipedes) along forest edge-to-interior gradients. Insect Conserv Diver. 2016;9(5):456-69. https://doi.org/10.1111/icad.12183.
    CrossRef
  22. de Souza Rezende R, Graça MAS, dos Santos AM, Medeiros AO, Santos PF, Nunes YR, et al. Organic matter dynamics in a tropical gallery forest in a grassland landscape. Biotropica. 2016;48(3):301-10. https://doi.org/10.1111/btp.12308.
    CrossRef
  23. Delang CO. Deforestation in northern Thailand: the result of Hmong farming practices or Thai development strategies? Soc Nat Resour. 2002;15(6):483-501. https://doi.org/10.1080/08941920290069137.
    CrossRef
  24. Dexter KG, Smart B, Baldauf C, Baker TR, Bessike Balinga MP, Brienen RJW, et al. Floristics and biogeography of vegetation in seasonally dry tropical regions. Int For Rev. 2015;17(S2):10-32.
    CrossRef
  25. Díaz-Aguilar I, Quideau SA. Trophic ecology of mesostigmatan and oribatid mites in harvested and control coniferous and deciduous stands of the boreal mixed wood forest determined using 15N stable isotopes. Soil Biol Biochem. 2013;67:147-54. https://doi.org/10.1016/j.soilbio.2013.08.019.
    CrossRef
  26. Didham RK. Altered leaf-litter decomposition rates in tropical forest fragments. Oecologia. 1998;116(3):397-406. https://doi.org/10.1007/s004420050603.
    Pubmed CrossRef
  27. Dossa GG, Paudel E, Cao K, Schaefer D, Harrison RD. Factors controlling bark decomposition and its role in wood decomposition in five tropical tree species. Sci Rep. 2016;6:34153. https://doi.org/10.1038/srep34153.
    Pubmed KoreaMed CrossRef
  28. Dumrongrojwatthana P. Impacts of forest disturbance on soil organic matter, soil nutrients and carbon sequestration in Nam Wa sub-watershed, Nan Province [MSc thesis]. Bangkok: Chulalongkorn University; 2004.
  29. Elmoghazy MME, Shawer SS. Relationship between soil diversity and inhabitant mites (Acari). Acarines. 2013;7(1):41-5.
    CrossRef
  30. Ewers RM, Didham RK. Confounding factors in the detection of species responses to habitat fragmentation. Biol Rev Camb Philos Soc. 2006;81(1):117-42. https://doi.org/10.1017/S1464793105006949.
    Pubmed CrossRef
  31. Ewers RM, Didham RK, Fahrig L, Ferraz G, Hector A, Holt RD, et al. A large-scale forest fragmentation experiment: the Stability of Altered Forest Ecosystems Project. Philos Trans R Soc Lond B Biol Sci. 2011;366(1582):3292-302. https://doi.org/10.1098/rstb.2011.0049.
    Pubmed KoreaMed CrossRef
  32. Feller IC, Whigham DF, McKee KL, Lovelock CE. Nitrogen limitation of growth and nutrient dynamics in a disturbed mangrove forest, Indian River Lagoon, Florida. Oecologia. 2003;134(3):405-14. https://doi.org/10.1007/s00442-002-1117-z.
    Pubmed CrossRef
  33. Ferguson SH. Does predation or moisture explain distance to edge distribution of soil arthropods? Am Midl Nat. 2004;152(1):75-87. https://doi.org/10.1674/0003-0031(2004)152[0075:DPOMED]2.0.CO;2.
    CrossRef
  34. Fujii S, Makita N, Mori AS, Takeda H. Plant species control and soil faunal involvement in the processes of above- and below-ground litter decomposition. Oikos. 2016;125(6):883-92. https://doi.org/10.1111/oik.02457.
    CrossRef
  35. Fuqiang S, Xiaoxu F, Ruiqing S. Review of mixed forest litter decomposition researches. Acta Ecol Sin. 2010;30(4):221-5. https://doi.org/10.1016/j.chnaes.2010.06.006.
    CrossRef
  36. García-Palacios P, Maestre FT, Kattge J, Wall DH. Climate and litter quality differently modulate the effects of soil fauna on litter decomposition across biomes. Ecol Lett. 2013;16(8):1045-53. https://doi.org/10.1111/ele.12137. Erratum in: Ecol Lett. 2013;16(11):1418.
    Pubmed KoreaMed CrossRef
  37. Gartner TB, Cardon ZG. Decomposition dynamics in mixed-species leaf litter. Oikos. 2004;104(2):230-46. https://doi.org/10.1111/j.0030-1299.2004.12738.x.
    CrossRef
  38. González G, Seastedt TR. Soil fauna and plant litter decomposition in tropical and subalpine forests. Ecology. 2001;82(4):955-64. https://doi.org/10.1890/0012-9658(2001)082[0955:SFAPLD]2.0.CO;2.
    CrossRef
  39. Guidi C, Frey B, Brunner I, Meusburger K, Vogel ME, Chen X, et al. Soil fauna drives vertical redistribution of soil organic carbon in a long-term irrigated dry pine forest. Glob Chang Biol. 2022;28(9):3145-60. https://doi.org/10.1111/gcb.16122.
    Pubmed KoreaMed CrossRef
  40. Hanpattanakit P, Chidthaisong A. Litter production and decomposition in dry dipterocarp forest and their responses to climatic factors. GMSARN Int J. 2012;6(2012):169-74.
  41. Hättenschwiler S, Tiunov AV, Scheu S. Biodiversity and litter decomposition in terrestrial ecosystems. Annu Rev Ecol Evol Syst. 2005;36:191-218. https://doi.org/10.1146/annurev.ecolsys.36.112904.151932.
    CrossRef
  42. Hättenschwiler S, Vitousek PM. The role of polyphenols in terrestrial ecosystem nutrient cycling. Trends Ecol Evol. 2000;15(6):238-43. https://doi.org/10.1016/s0169-5347(00)01861-9.
    Pubmed CrossRef
  43. Ishida A, Yamaji K, Nakano T, Ladpala P, Popradit A, Yoshimura K, et al. Comparative physiology of canopy tree leaves in evergreen and deciduous forests in lowland Thailand. Sci Data. 2023;10(1):601. https://doi.org/10.1038/s41597-023-02468-6.
    Pubmed KoreaMed CrossRef
  44. Khaing TT, Pasion BO, Lapuz RS, Tomlinson KW. Determinants of composition, diversity and structure in a seasonally dry forest in Myanmar. Glob Ecol Conserv. 2019;19:e00669. https://doi.org/10.1016/j.gecco.2019.e00669.
    CrossRef
  45. Knorr M, Frey SD, Curtis PS. Nitrogen additions and litter decomposition: a meta-analysis. Ecology. 2005;86(12):3252-7.
    CrossRef
  46. Koné AW, Edoukou EF, Gonnety JT, N'Dri AN, Assémien LF, Angui PK, et al. Can the shrub Chromolaena odorata (Asteraceae) be considered as improving soil biology and plant nutrient availability? Agroforest Syst. 2012;85:233-45. https://doi.org/10.1007/s10457-012-9497-5.
    CrossRef
  47. Koné AW, Kassi SPAY, Koffi BY, Masse D, Maïga AA, Tondoh JE, et al. Chromolaena odorata (L.) K&R (Asteraceae) invasion effects on soil microbial biomass and activities in a forest-savanna mosaic. Catena. 2021;207:105619. https://doi.org/10.1016/j.catena.2021.105619.
    CrossRef
  48. Krishna M, Mohan M. Litter decomposition in forest ecosystems: a review. Energ Ecol Environ. 2017;2:236-49. https://doi.org/10.1007/s40974-017-0064-9.
    CrossRef
  49. Lacasella F, Gratton C, De Felici S, Isaia M, Zapparoli M, Marta S, et al. Asymmetrical responses of forest and "beyond edge" arthropod communities across a forest-grassland ecotone. Biodivers Conserv. 2015;24:447-65. https://doi.org/10.1007/s10531-014-0825-0.
    CrossRef
  50. Laurance WF, Camargo JLC, Luizão RCC, Laurance SG, Pimm SL, Bruna EM, et al. The fate of Amazonian forest fragments: a 32-year investigation. Biol Conserv. 2011;144(1):56-67. https://doi.org/10.1016/j.biocon.2010.09.021.
    CrossRef
  51. Li A, Fan Y, Chen S, Song H, Lin C, Yang Y. Soil warming did not enhance leaf litter decomposition in two subtropical forests. Soil Biol Biochem. 2022;170:108716. https://doi.org/10.1016/j.soilbio.2022.108716.
    CrossRef
  52. Manu M. The influence of some environmental factors on the species diversity of the predator mites (Acari: Mesostigmata) from natural forest ecosystems of Bucegi Massif (Romania). Trav Mus Natl Hist Nat "Grigore Antipa" 2011;LIV(1):9-20. https://doi.org/10.2478/v10191-011-0001-7.
    CrossRef
  53. Matsumoto T, Abe T. The role of termites in an equatorial rain forest ecosystem of West Malaysia: II. Leaf litter consumption on the forest floor. Oecologia. 1979;38(3):261-74. https://doi.org/10.1007/BF00345187.
    Pubmed CrossRef
  54. Mboukou-Kimbatsa I, Bernhard-Reversat F, Loumeto JJ, Ngao J, Lavelle P. Understory vegetation, soil structure and soil invertebrates in Congolese eucalypt plantations, with special reference to the invasive plant Chromolaena odorata and earthworm populations. Eur J Soil Biol. 2007;43(1):48-56. https://doi.org/10.1016/j.ejsobi.2006.05.002.
    CrossRef
  55. Meyer S, Rusterholz HP, Salamon JA, Baur B. Leaf litter decomposition and litter fauna in urban forests: effect of the degree of urbanisation and forest size. Pedobiologia. 2020;78:150609. https://doi.org/10.1016/j.pedobi.2019.150609.
    CrossRef
  56. Mondal N, Sukumar R. Characterising weather patterns associated with fire in a seasonally dry tropical forest in southern India. Int J Wildland Fire. 2013;23(2):196-201. https://doi.org/10.1071/WF13002.
    CrossRef
  57. Moreno ML, Bernaschini ML, Pérez-Harguindeguy N, Lomba A, Valladares G. Chaco forest fragmentation effects on leaf litter decomposition are not explained by changes in litter fauna. Austral Ecol. 2020;45(1):27-34. https://doi.org/10.1111/aec.12824.
    CrossRef
  58. Moreno ML, Bernaschini ML, Pérez-Harguindeguy N, Valladares G. Area and edge effects on leaf-litter decomposition in a fragmented subtropical dry forest. Acta Oecol. 2014;60:26-9. https://doi.org/10.1016/j.actao.2014.07.002.
    CrossRef
  59. Myo KK, Thwin S, Khaing N. Floristic composition, structure and soil properties of mixed deciduous forest and deciduous dipterocarp forest: case study in Madan Watershed, Myanmar. Am J Plant Sci. 2016;7(2):279-87. https://doi.org/10.4236/ajps.2016.72027.
    CrossRef
  60. Njoroge DM, Chen SC, Zuo J, Dossa GGO, Cornelissen JHC. Soil fauna accelerate litter mixture decomposition globally, especially in dry environments. J Ecol. 2022;110(3):659-72. https://doi.org/10.1111/1365-2745.13829.
    CrossRef
  61. Njoroge DM, Dossa GGO, Ye L, Lin X, Schaefer D, Tomlinson K, et al. Fauna access outweighs litter mixture effect during leaf litter decomposition. Sci Total Environ. 2023;860:160190. https://doi.org/10.1016/j.scitotenv.2022.160190.
    Pubmed CrossRef
  62. Norgrove L, Hauser S, Weise SF. Response of Chromolaena odorata to timber tree densities in an agrisilvicultural system in Cameroon: aboveground biomass, residue decomposition and nutrient release. Agric Ecosyst Environ. 2000;81(3):191-207. https://doi.org/10.1016/S0167-8809(00)00171-7.
    CrossRef
  63. Ochoa-Hueso R, Delgado-Baquerizo M, King PTA, Benham M, Arca V, Power SA. Ecosystem type and resource quality are more important than global change drivers in regulating early stages of litter decomposition. Soil Biol Biochem. 2019;129:144-52. https://doi.org/10.1016/j.soilbio.2018.11.009.
    CrossRef
  64. Okeke AI, Omaliko CPE. Leaf litter decomposition and carbon dioxide evolution of some agroforestry fallow species in southern Nigeria. For Ecol Manage. 1992;50(1-2):103-16. https://doi.org/10.1016/0378-1127(92)90317-3.
    CrossRef
  65. Olson JS. Energy storage and the balance of producers and decomposers in ecological systems. Ecology. 1963;44(2):322-31. https://doi.org/10.2307/1932179.
    CrossRef
  66. Ong CSP, Juan JC, Yule CM. The contribution of leaching to nutrient release from leaf litter of two emergent tree species in a Malaysian tropical peat swamp forest. Hydrobiologia. 2017;794:125-37. https://doi.org/10.1007/s10750-017-3086-6.
    CrossRef
  67. Osono T. Functional diversity of ligninolytic fungi associated with leaf litter decomposition. Ecol Res. 2020;35(1):30-43. https://doi.org/10.1111/1440-1703.12063.
    CrossRef
  68. Pamminger T, Bottoms M, Cunningham H, Ellis S, Kabouw P, Kimmel S, et al. Investigating the role of soil mesofauna abundance and biodiversity for organic matter breakdown in arable fields. Integr Environ Assess Manag. 2022;18(5):1423-33. https://doi.org/10.1002/ieam.4563.
    Pubmed KoreaMed CrossRef
  69. Paudel E, Dossa GGO, de Blécourt M, Beckschäfer P, Xu J, Harrison RD. Quantifying the factors affecting leaf litter decomposition across a tropical forest disturbance gradient. Ecosphere. 2015;6(12):1-20. https://doi.org/10.1890/ES15-00112.1.
    CrossRef
  70. Peguero G, Sardans J, Asensio D, Fernández-Martínez M, Gargallo-Garriga A, Grau O, et al. Nutrient scarcity strengthens soil fauna control over leaf litter decomposition in tropical rainforests. Proc R Soc B. 2019;286(1910):20191300. https://doi.org/10.1098/rspb.2019.1300.
    Pubmed KoreaMed CrossRef
  71. R Core Team. R: a language and environment for statistical computing. Vienna: R Foundation for Statistical Computing; 2022.
  72. Robbins CJ, Matthaeus WJ, Cook SC, Housley LM, Robison SE, Garbarino MA, et al. Leaf litter identity alters the timing of lotic nutrient dynamics. Freshw Biol. 2019;64(12):2247-59. https://doi.org/10.1111/fwb.13410.
    CrossRef
  73. Sabatté ML, Massobrio MJ, Cassani MT, Momo FR. Macro and mesofauna soil food webs in two temperate grasslands: responses to forestation with Eucalyptus. Heliyon. 2021;7(1):e05869. https://doi.org/10.1016/j.heliyon.2020.e05869.
    Pubmed KoreaMed CrossRef
  74. Salles LFP, Christianini AV, Oliveira PS. Dirt roads and fire breaks produce no edge effects on litter-dwelling arthropods in a tropical dry-forest: a case study. J Insect Conserv. 2018;22:647-57. https://doi.org/10.1007/s10841-018-0091-7.
    CrossRef
  75. Santonja M, Fernandez C, Proffit M, Gers C, Gauquelin T, Reiter IM, et al. Plant litter mixture partly mitigates the negative effects of extended drought on soil biota and litter decomposition in a Mediterranean oak forest. J Ecol. 2017;105(3):801-15. https://doi.org/10.1111/1365-2745.12711.
    CrossRef
  76. Sarjubala Devi A, Yadava PS. Wood and leaf litter decomposition of Dipterocarpus tuberculatus Roxb. in a tropical deciduous forest of Manipur, Northeast India. Curr Sci. 2007;93(2):243-6.
  77. Seidelmann KN, Scherer-Lorenzen M, Niklaus PA. Direct vs. microclimate-driven effects of tree species diversity on litter decomposition in young subtropical forest stands. PLoS One. 2016;11(8):e0160569. https://doi.org/10.1371/journal.pone.0160569.
    Pubmed KoreaMed CrossRef
  78. Sharma J, Singh R, Garai S, Rahaman SM, Khatun M, Ranjan A, et al. Climate change and dispersion dynamics of the invasive plant species Chromolaena odorata and Lantana camara in parts of the central and eastern India. Ecol Inform. 2022;72:101824. https://doi.org/10.1016/j.ecoinf.2022.101824.
    CrossRef
  79. Sodhi NS, Koh LP, Clements R, Wanger TC, Hill JK, Hamer KC, et al. Conserving Southeast Asian forest biodiversity in human-modified landscapes. Biol Conserv. 2010;143(10):2375-84. https://doi.org/10.1016/j.biocon.2009.12.029.
    CrossRef
  80. Steinwandter M, Schlick-Steiner BC, Steiner FM, Seeber J. One plus one is greater than two: mixing litter types accelerates decomposition of low-quality alpine dwarf shrub litter. Plant Soil. 2019;438:405-19. https://doi.org/10.1007/s11104-019-03991-5.
    CrossRef
  81. Szigeti N, Berényi Üveges J, Berki I, Vityi A. Grassy-floral soil covering as a tool for increasing herbaceous diversity in agroforestry. J Cent Eur Agric. 2022;23(4):898-908. https://doi.org/10.5513/JCEA01/23.4.3672.
    CrossRef
  82. Tedersoo L, Bahram M, Cajthaml T, Põlme S, Hiiesalu I, Anslan S, et al. Tree diversity and species identity effects on soil fungi, protists and animals are context dependent. ISME J. 2016;10(2):346-62. https://doi.org/10.1038/ismej.2015.116.
    Pubmed KoreaMed CrossRef
  83. Tie L, Wei S, Peñuelas J, Sardans J, Liu X, Zhou S, et al. N and P combined addition accelerates the release of litter C, N, and most metal nutrients in a N-rich subtropical forest. Sci Total Environ. 2023;881:163491. https://doi.org/10.1016/j.scitotenv.2023.163491.
    Pubmed CrossRef
  84. Walkley A, Black IA. An examination of the Degtjareff method for determining soil organic matter, and a proposed modification of the chromic acid titration method. Soil Sci. 1934;37(1):29-38.
    CrossRef
  85. Wang D, Yuan F, Xie W, Zuo J, Zhou H. Effects of leaf size and defensive traits on the contribution of soil fauna to litter decomposition. Forests. 2024;15(3):481. https://doi.org/10.3390/f15030481.
    CrossRef
  86. Wang Y, Wu J, Ma D, Pu H, Yin Q, Chen W. Study on macro-meso mechanical properties of cemented tailings backfill with high fly ash content. Environ Sci Pollut Res Int. 2023;30(2):2904-17. https://doi.org/10.1007/s11356-022-22436-8.
    Pubmed CrossRef
  87. Wanthongchai K, Bauhus J, Goldammer JG. Nutrient losses through prescribed burning of aboveground litter and understorey in dry dipterocarp forests of different fire history. Catena. 2008;74(3):321-32. https://doi.org/10.1016/j.catena.2008.01.003.
    CrossRef
  88. Wei H, Xu J, Quan G, Zhang J, Qin Z. Invasion effects of Chromolaena odorata on soil carbon and nitrogen fractions in a tropical savanna. Ecosphere. 2017;8(5):e01831. https://doi.org/10.1002/ecs2.1831.
    CrossRef
  89. Xin WD, Yin XQ, Song B. Contribution of soil fauna to litter decomposition in Songnen sandy lands in northeastern China. J Arid Environ. 2012;77:90-5. https://doi.org/10.1016/j.jaridenv.2011.10.001.
    CrossRef
  90. Yang K, Zhu J, Zhang W, Zhang Q, Lu D, Zhang Y, et al. Litter decomposition and nutrient release from monospecific and mixed litters: comparisons of litter quality, fauna and decomposition site effects. J Ecol. 2022;110(7):1673-86. https://doi.org/10.1111/1365-2745.13902.
    CrossRef
  91. Yang L, Han T, Fu Y. Effect of heat treatment and wax impregnation on dimensional stability of Pterocarpus macrocarpus wood. Wood Res. 2020;65(6):963-74. https://doi.org/10.37763/wr.1336-4561/65.6.963974.
    CrossRef
  92. Yang X, Chen J. Plant litter quality influences the contribution of soil fauna to litter decomposition in humid tropical forests, southwestern China. Soil Biol Biochem. 2009;41(5):910-8. https://doi.org/10.1016/j.soilbio.2008.12.028.
    CrossRef
  93. Zeileis A, Hothorn T. Diagnostic checking in regression relationships. R News. 2002;2(3):7-10.
  94. Zhang Q, Zak JC. Effects of gap size on litter decomposition and microbial activity in a subtropical forest. Ecology. 1995;76(7):2196-204. https://doi.org/10.2307/1941693.
    CrossRef
  95. Zheng X, Tao Y, Wang Z, Kou X, Wang H, Wang S, et al. Land-use types influence the community composition of soil mesofauna in the coastal zones of Bohai Bay, China. Diversity. 2022;14(12):1035. https://doi.org/10.3390/d14121035.
    CrossRef

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