Soil Mechanical Composition and Moisture Enhanced the Improvement of Mealworm Casting Amendment on Different Land Use Soils

Soil Mechanical Composition and Moisture Enhanced the Improvement of Mealworm Casting Amendment on Different Land Use Soils | Research Square window.SnipcartSettings = { analytics: { enabled: false } }; (function() { var accessVector = localStorage.getItem('access_vector') || ''; window.dataLayer = window.dataLayer || []; if (accessVector) { window.dataLayer.push({ user: { profile: { profileInfo: { snid: accessVector } } } }); } })(); (function(w,d,s,l,i){w[l]=w[l]||[];w[l].push({'gtm.start':new Date().getTime(),event:'gtm.js'});var f=d.getElementsByTagName(s)[0],j=d.createElement(s),dl=l!='dataLayer'?'&l='+l:'';j.async=true;j.src='https://www.googletagmanager.com/gtm.js?id='+i+dl;f.parentNode.insertBefore(j,f);})(window,document,'script','dataLayer','GTM-K279D39R'); Browse Preprints In Review Journals COVID-19 Preprints AJE Video Bytes Research Tools Research Promotion AJE Professional Editing AJE Rubriq About Preprint Platform In Review Editorial Policies Our Team Advisory Board Help Center Sign In Submit a Preprint Cite Share Download PDF Research Article Soil Mechanical Composition and Moisture Enhanced the Improvement of Mealworm Casting Amendment on Different Land Use Soils Zhi-Shen Hong, Cai-Yi Zhu, Xiao-Qing Li, Ruofei Wang, Zeyu Li, and 4 more This is a preprint; it has not been peer reviewed by a journal. https://doi.org/ 10.21203/rs.3.rs-8998912/v1 This work is licensed under a CC BY 4.0 License Status: Posted Version 1 posted You are reading this latest preprint version Abstract To investigate the effects of mealworm casting amendment on different land-use soils and find out the optimal soil moisture content for application, three distinct soils of different soil texture were selected for soil incubation experiment, one of which was set to two moisture levels. Soil samples were collected on days 14 and 28 of the incubation. Research findings indicate that soil mealworm casting amendemnt stimulates soil carbon and nitrogen cycling, facilitating the conversion of nutrients into forms readily absorbed by plants. The most pronounced effect is observed in soil ammonium nitrogen content, with an average increase of 620.7%; the average increases in soluble carbon and soluble organic carbon were 80.7% and 92.5%, respectively. Adding insect manure significantly increased the abundance of microorganisms involved in the soil carbon cycling, such as Proteobacteria and Bacteroidetes. The mealworm manure also increased beneficial taxa such as Chitinophaga and suppressed disease-associated microorganisms, thereby improving the soil microbial community structure. Moisture conditions and soil mechanical composition were key factors influencing the effectiveness of mealworm castings. The optimal moisture content for mealworm castings application is 60% of the field capacity, while higher moisture (80% of the field capacity) inhibits casting mineralization and reduces soil nitrate nitrogen content by 73.2% at day 28. Suitable moisture contents promoted mineralization of Tenebrio molitor castings, whereas excessive clay content constrained both mineralization and the frass-induced increase in soil pH. Tenebrio molitor castings organic fertilizer soil amendment soil microbial community structure amplicon sequencing Figures Figure 1 Figure 2 Figure 3 Figure 4 Figure 5 Figure 6 Figure 7 Figure 8 Figure 9 1 Introduction Tenebrio molitor is a high-quality protein source and can be used as a premium feed additive for livestock or as a protein supplement in certain foods. Owing to its considerable economic value, it is widely cultivated on a large scale. As a holometabolous insect, the yellow mealworm is primarily commercialized at the larval stage before pupation. Large-scale larval rearing generates substantial quantities of mealworm castings (frass). Previous reports indicate that the ratio of Tenebrio molitor castings to dry Tenebrio molitor biomass is approximately 8.8:1. In 2021, the European Food Safety Authority (EFSA) formally authorized Tenebrio molitor as a novel food for human consumption. In China, Tenebrio molitor production reached 61,600 tons in 2023, while demand was 53,400 tons, and both figures continue to increase annually. Therefore, the efficient and scientifically sound utilization of mealworm castings in agricultural production to maximize their value remains an urgent issue that needs to be addressed. Mealworm castings are byproducts of yellow mealworm farming. They are typically described as dry, odorless, and sand-like, with a fine granular structure, and are easy to store and transport (Gao Yan, 2012; Shen, X. K. et al ., 2009). Studies have shown that mealworm castings are rich in organic matter and essential plant nutrients, including nitrogen, phosphorus, and potassium, and also contain insect-derived metabolites such as chitin and antimicrobial peptides. Compared with traditional livestock manure, mealworm castings have a more balanced nitrogen-to-phosphorus ratio and lower heavy metal contents, making them more environmentally friendly. In addition, they can improve soil structure, increase water-holding capacity, and enhance soil fertility (He et al., 2021 ; Zunzunegui et al., 2024 ). Currently, few studies have examined the differential effects of mealworm castings across soils from different land-use types. Variation in the effects of organic fertilizers among land-use types represents a complex ecological issue shaped by multiple factors, including soil physicochemical properties, microbial communities, vegetation, and management practices (Li et al ., 2021). Numerous studies have shown that organic fertilizers can exert markedly different effects on soil fertility, nutrient cycling, and ecosystem functioning across croplands, grasslands, and forest lands (Fang et al ., 2013; Tang et al., 2020 ; Zajicova et al ., 2019). Soil texture—specifically the proportions of sand, silt, and clay—regulates the effectiveness of organic fertilizers and the extent of fertility improvement by influencing soil physical structure, water-holding capacity, nutrient adsorption and release, and microbial activity (Adeniji, A. et al ., 2021). In addition, the bioavailability of soil organic carbon (SOC) is closely linked to microbial activity and is mediated by particle-size fractionation and soil morphology (Xiao, S. et al., 2024 ). Therefore, optimizing mealworm casting application strategies according to soil texture is essential for enhancing agricultural productivity, improving soil health, and promoting sustainable agriculture. Croplands, as the primary land-use type receiving organic fertilizer inputs, are also expected to be a major target for large-scale application of mealworm castings. Moderate soil moisture (typically close to field capacity) promotes the mineralization of organic matter, thereby facilitating the release of nutrients such as nitrogen, phosphorus, and potassium from organic fertilizers for crop uptake (Du, T et al ., 2022). At the same time, soil moisture is a key environmental factor shaping microbial communities; changes in moisture can alter microbial community composition and diversity (Xing, Y et al., 2024 ). Moreover, organic fertilizer inputs and moisture conditions can jointly influence the functional diversity of key soil microorganisms. Organic fertilizers can exert broad positive effects on soil environments and crop yields by improving soil physicochemical properties and reshaping microbial communities (Ma, G. et al., 2023 ; Xing, Y et al., 2024 ). However, their effectiveness varies across land-use practices and is largely determined by initial soil fertility, soil texture, management practices (e.g., water management), and ecosystem characteristics (Bajgai, Y. et al ., 2023; Li, Z. et al ., 2024). In this study, we investigated the effects of adding mealworm castings at 1% (w/w) to cropland soils representing three land-use types. We further analyzed how soil moisture conditions modulate the impacts of mealworm castings on soil physicochemical properties and microbial communities, thereby providing a theoretical basis for the scientific utilization of mealworm castings. 2 Materials and methods 2.1 Test soil and mealworm castings To fugure out the effect of mealworm castings on soils, three soils of cropland, forest land and grassland with different soil texture were selected for soil incubation experiment. The cropland soil was sampled from Binzhou City, Shandong Province (37.3025N,117.366E), with a land use type of irrigated farmland; the forest land soil was collected from Guangzhou City, Guangdong Province (23.930N,113.2120E), classified as forest land; the grassland soil was collected from Guangzhou City, Guangdong Province (23.936N,113.2043E), classified as other grassland. The collected soil samples were coarsely ground, sieved through a 2 mm mesh, and naturally air-dried. Field water-holding capacity, particle size distribution, and moisture content were then measured. Land use types were classified according to GB/T 21010 − 2007. Basic physicochemical properties of soil and mealworm castings are detailed in Tables 1 . Based on the International Soil Classification Triangle, all three soil textures were classified as sandy loam. The mealworm castings used in this experiment were obtained from Tenebrio molitor larvae reared with wheat bran in the laboratory. After collecting, the castings were sieved through a 60-mesh screen. Table 1 .Basic Physical and Chemical Properties of Test Soils Soil sample M DL LN FT pH 6.40 ± 0.03 7.69 ± 0.00 7.63 ± 0.00 4.86 ± 0.00 TC/(g·kg − 1 ) 395.0 ± 2.4 21.1 ± 0.1 6.3 ± 0.1 23.8 ± 0.1 TN/(g·kg − 1 ) 58.20 ± 0.43 1.67 ± 0.06 0.67 ± 0.01 1.37 ± 0.30 Gravel content % - 65.8 ± 0.0 72.1 ± 0.0 76.7 ± 0.0 Particle content % - 25.2 ± 0.0 26.9 ± 0.0 14.1 ± 0.0 Silt content % - 9.03 ± 0.00 1.01 ± 0.00 9.21 ± 0.00 Field capacity % - 36.8 ± 0.0 48.3 ± 0.0 37.8 ± 0.0 Air dried soil Moisture content % - 1.87 ± 0.00 1.96 ± 0.00 1.12 ± 0.00 TSC/(mg·kg − 1 ) 17820.7 ± 1979.6 138.1 ± 16.3 28.1 ± 2.6 135.0 ± 20.6 TSOC/(mg·kg − 1 ) 16496.6 ± 1431.6 111.6 ± 5.3 26.1 ± 0.7 133.0 ± 9.5 AvbN/(mg·kg − 1 ) - 78.2 ± 1.7 18.4 ± 0.6 102.3 ± 0.8 NH 4 + -N/(mg·kg − 1 ) - 9.85 ± 1.14 3.45 ± 0.64 26.77 ± 4.24 NO 3 − -N/(mg·kg − 1 ) - 50.97 ± 10.68 4.71 ± 1.57 19.23 ± 4.84 AvbP/(mg·kg − 1 ) - 70.49 ± 9.20 5.96 ± 3.43 4.71 ± 0.91 AvbK/(mg·kg − 1 ) - 113.2 ± 0.6 31.9 ± 0.0 29.4 ± 3.6 pH, soil acidity/alkalinity (soil-water ratio 1:2.5); TN, total nitrogen; TC, total carbon; TSC, soluble carbon; TSOC, soluble organic carbon; AvbN, alkali-hydrolyzable nitrogen; NH 4 + -N, ammonium; NO 3 − -N, nitrogen; AvbP, available phosphorus; AvbK, available potassium. Table values are presented as mean ± standard deviation. M Tenebrio molitor castings; DL denotes croplands; LN denotes grassland; FT denotes forest land. 2.2 Experimental Design Eight treatments were established in this experiment (see Table 2 for details), with 3 replicates per treatment. The specific experimental procedure is as follows: Each beaker contained 100 g of dried soil (converted to air-dried soil based on moisture content). Samples were divided into separate 200 ml beakers according to treatment. Neatly arrange the beakers containing the divided samples in a plastic box. Place water at the bottom of the box for humidity maintenance, cover the box, and place it in a constant-temperature incubator. Incubate at 26°C. Every three days, add an appropriate amount of ultrapure water to each treatment sample to restore it to its original weight. Destructive sampling was performed on days 14 and 28, followed by post-treatment preservation. A portion of samples was air-dried for storage; another portion was stored at 4°C for basic physicochemical parameter analysis; and a third portion was preserved at -80°C for microbial parameter analysis. Table 2 Basic Information for Each Treatment Processing Number Land Use Type Soil Type Code Processing Method D6M Cropland DL Add 1% mealworm castings (based on 1% of the dry weight of the soil sample) at 60% of the field capacity. D6 Cropland DL 60% of the field capacity D8M Cropland DL Add 1% mealworm castings (based on 1% of the dry weight of the soil sample) at 80% of the field capacity. D8 Cropland DL 80% of the field capacity L6M Grassland LN Add 1% mealworm castings (based on 1% of the dry weight of the soil sample) at 60% of the field capacity. L6 Grassland LN 60% of the field capacity F6M Forest land FT Add 1% mealworm castings (based on 1% of the dry weight of the soil sample) at 60% of the field capacity. F6 Forest land FT 60% of the field capacity 2.3 Soil Physicochemical Properties Testing The determination of soil physicochemical parameters followed Bao Shidan's Soil Agrochemical Analysis (China Agricultural Press, 2000). Soil pH (soil : water = 1 : 2.5) was measured using the potentiometric method; soil electrical conductivity was determined under extraction conditions consistent with soil pH, followed by measurement with a conductivity meter; soil available phosphorus content was determined using the ammonium fluoride extraction method; soil available potassium was determined by flame photometry; soil total nitrogen and total carbon were measured using a fully automated elemental analyzer (Manufacturer: Elementar, Model: Vario Micro cube); soil soluble organic carbon and soluble total nitrogen were determined by water extraction using a total organic carbon analyzer (Manufacturer: Elementar, Model: Vario TOC); Soil ammonium nitrogen was determined using the indophenol blue colorimetric method; Soil nitrate nitrogen was determined using the potassium chloride extraction method; Alkali-hydrolyzable nitrogen was determined using boric acid absorption titration. 2.4 Soil Microbial Sequencing Microbial community analysis for this experiment was conducted by Beijing Novogene Co., Ltd., which performed extraction and sequencing using the Illumina NovaSeq PE250 platform. Fungi were sequenced using ITS amplicon sequencing, targeting the ITS1-1F region with primer sequences: ITS5-1737 F (5’-CTTGGTCATTTAGAGGAAGTAA-3’) and ITS2-2043 R (5’-GCTGCGTTCTTCATCGATGC-3’). Bacteria were sequenced using the 16S rDNA amplicon, with the amplification region set to 16S V3-V4. Primer sequences were: 515 F (5’-CCTAYGGGRBGCASCAG-3’) and 806 R (5’-GGACTACNNGGGTATCTAAT-3’). 1.5 Data Analysis and Statistical Methods Data analysis and processing were performed using Microsoft Office Excel 2013 (Microsoft Corporation, USA), SPSS 18.0 (IBM Corporation), and RStudio (RStudio Company). Alpha diversity and Beta diversity analyses were conducted using the R software version 4.2.3. The raw sequencing data (Raw Data) contained a certain proportion of interfering data (Dirty Data). To ensure that the results of the information analysis were more accurate and reliable, the raw data were first merged and filtered to obtain effective data (Clean Data). Then, the effective data were denoised using DADA2 or deblur (Li M et al., 2020 ), thereby yielding the final ASVs. For the obtained ASVs (Callahan BJ et al ., 2017), on the one hand, the representative sequence of each ASV was subjected to taxonomic annotation to obtain the corresponding species information and species-based abundance distribution. The classify-sklearn algorithm in QIIME2 (Bokulich NA et al., 2018 ; Bolyen E et al ., 2019) was used to annotate each ASV using a pre-trained Naive Bayes classifier. The annotation database was Silva 138.1. 3 Results Analysis 3.1 Effects of Tenebrio molitor Castings on Soil Physicochemical Properties As shown in Table 3 , compared with the blank control, the addition of mealworm castings significantly increased the content of soil total nitrogen (TN), soil soluble carbon (TSC), soil soluble organic carbon (TSOC), alkali-hydrolyzable nitrogen (AvbN), ammonium nitrogen (NH 4 + -N), and soil nitrate nitrogen (NO 3 − -N) in both forest (FT) and grassland (LN) soils. The increases ranged from 15.0% to 31.6%, 37.2% to 171.1%, 41.5% to 178.0%, 73.9% to 203.8%, 184.0% to 620.7%, and 31.5% to 581.9%, respectively. The addition of mealworm castings significantly increased the pH value of forest land by 0.3–0.5 units. For grassland, the pH value decreased significantly by 0.2 units on day 28. Compared to the blank control, insect dung addition also significantly increased available potassium (AvbK) and available phosphorus (AvbP) in forest land (FT), grassland (LN), and cropland (DL) (P < 0.05), with increases ranging from 116% to 672% for AvbK and from 23.7% to 726.7% for AvbP. Among different cropland-capacity moisture treatments, adding mealworm castings had no significant effect on soil soluble carbon content in the 60% cropland-capacity treatment (DL6). However, it significantly increased soluble carbon content in the 80% cropland-capacity treatment (DL8) by 56.3% on day 28. Compared to the blank control, the DL6 treatment with added mealworm castings showed a significant increase in soil soluble organic carbon at day 14, while the DL8 treatment exhibited a significant increase only at day 28, with increases of 49.2% and 56.3%, respectively. Compared to the blank control, adding mealworm castings significantly increased total soil nitrogen in the DL6 treatment, with increases ranging from 11.3% to 11.7%. In the DL8 treatment, adding mealworm castings significantly reduced soil nitrate nitrogen content compared to the blank, with a decrease ranging from 73.2% to 87.4%. In DL treatments, ammonium nitrogen content significantly increased by 128%–176% at day 28. In the DL8 treatment, soil ammonium nitrogen content increased significantly on day 14, with an increase of 152.3%. DL6 treatment with mealworm castings significantly lowered pH by 0.08–0.16 units, while DL8 treatment with mealworm castings significantly raised pH by 0.08–0.14 units. As shown in Fig. 1 , PC1 explained 50.34% of the variance, PC2 explained 24.98%, and the cumulative explanation reached 75.32%. The treatments incorporating mealworm castings showed distinct differentiation from the blank control and original soil samples. Compared to the blank control, the addition of mealworm castings resulted in a clear differentiation along the PC1 axis overall. This indicates that adding mealworm castings significantly increased soil alkaline-hydrolyzable nitrogen content, soil soluble nitrogen content, soil soluble carbon content, soil total carbon content, soil total nitrogen content, ammonium nitrogen content, nitrate nitrogen content, available phosphorus content, and available potassium content. The treatment with added mealworm castings showed distinct differentiation on the PC2 axis. This indicates that over time, the addition of mealworm castings does not affect soil total carbon content, has a minor impact on total nitrogen content, but significantly influences pH and ammonium nitrogen content. Among treatments with different moisture contents, the treatment at 60% field capacity showed more pronounced differentiation on the PC1 axis compared to the treatment at 80% field capacity. Table 3 Effects of Different Treatments on Soil Physicochemical Properties sample pH TN % TC % TSC /(mg·kg − 1 ) TSOC /(mg·kg − 1 ) AvbN /(mg·kg − 1 ) NH 4 + -N /(mg·kg − 1 ) NO 3 − -N /(mg·kg − 1 ) AvbP /(mg·kg − 1 ) AvbK /(mg·kg − 1 ) d14D6M 7.35 ± 0.02c 1.90 ± 0.10a 22.33 ± 0.7a 134.2 ± 36.5a 102.4 ± 11.5a 84.3 ± 3.0a 5.66 ± 0.94bc 39.56 ± 5.85a 89.2 ± 9.0ab 280.6 ± 7.1a d14D6 7.44 ± 0.05b 1.70 ± 0.00b 20.83 ± 0.35b 89.9 ± 1.3a 65.9 ± 3.2b 73.9 ± 3.3b 6.37 ± 1.31b 42.4 ± 9.64a 61.5 ± 9.8b 123.7 ± 7.2c d14D8M 7.61 ± 0.03a 1.67 ± 0.06a 23.50 ± 0.70a 98.4 ± 16.6ab 60.5 ± 2.3bc 84.9 ± 5.1a 12.78 ± 1.32b 4.25 ± 0.22c 106.0 ± 9.0a 260.1 ± 2.2a d14D8 7.47 ± 0.02b 1.63 ± 0.06a 20.83 ± 0.35c 112.3 ± 5.9a 71 ± 13.5b 74.9 ± 8.2a 5.07 ± 0.61c 33.69 ± 6.52a 68.8 ± 2.1b 116.9 ± 0.6b d14L6M 7.21 ± 0.05a 0.83 ± 0.12a 8.40 ± 0.60a 41.3 ± 4.7a 38 ± 1.5a 65.8 ± 6.5a 28.19 ± 0.99a 14.79 ± 0.78b 10.9 ± 0.8a 205.8 ± 9.7a d14L6 7.23 ± 0.04a 0.67 ± 0.06b 6.63 ± 0.61b 15.2 ± 2.2c 13.7 ± 0.5c 21.7 ± 0.3c 3.91 ± 0.57c 2.81 ± 0.08c 1.3 ± 0.4b 33.4 ± 1.6b d14F6M 5.52 ± 0.03a 1.53 ± 0.06a 21.17 ± 0.76ab 149.2 ± 17.7a 144.8 ± 20.2a 186.0 ± 7.0a 42.91 ± 3.79a 23.04 ± 6.28b 24.2 ± 2.6b 196.9 ± 7.8a d14F6 4.99 ± 0.06b 1.33 ± 0.06b 20.57 ± 0.84b 108.7 ± 23.6b 102.3 ± 18b 99.8 ± 7.1c 8.41 ± 0.90c 6.14 ± 1.61c 7.5 ± 1.7c 25.1 ± 1.2b d28D6M 7.41 ± 0.04bc 1.97 ± 0.06a 22.13 ± 0.21a 107.8 ± 24.5a 66 ± 9.6b 76.8 ± 6.6ab 10.03 ± 2.10a 45.94 ± 6.83a 99.2 ± 24.3a 254.6 ± 0.5b d28D6 7.57 ± 0.06a 1.77 ± 0.06b 21.13 ± 0.57b 107.3 ± 24.3a 65.7 ± 9.5b 76.7 ± 1.4b 3.63 ± 0.79c 38.94 ± 3.57a 80.2 ± 1.0ab 117.3 ± 1.2c d28D8M 7.58 ± 0.04a 1.67 ± 0.06a 22.37 ± 0.65ab 123.2 ± 20.7a 89.4 ± 2.5a 85.9 ± 4.5a 24.15 ± 1.42a 6.45 ± 1.37c 110.7 ± 4.3a 260.4 ± 0.6a d28D8 7.50 ± 0.01b 1.67 ± 0.06a 21.47 ± 0.91bc 78.8 ± 2.3b 52.5 ± 8.6c 73.8 ± 8.4a 10.58 ± 2.20b 24.03 ± 2.99b 78.9 ± 6.8b 117.3 ± 1.8b d28L6M 7.04 ± 0.04b 0.83 ± 0.06a 8.80 ± 0.70a 34.5 ± 0.8b 33.1 ± 1.2b 53.8 ± 1.7b 13.55 ± 1.41b 18.63 ± 1.33a 12.7 ± 2.3a 198.3 ± 4a d28L6 7.24 ± 0.03a 0.63 ± 0.06b 6.83 ± 0.31b 14.5 ± 2.1c 13.4 ± 0.7c 24.1 ± 2.2c 4.77 ± 0.75c 2.73 ± 0.43c 3.7 ± 2.2b 30.4 ± 1.1b d28F6M 4.98 ± 0.02b 1.57 ± 0.15a 21.17 ± 0.61ab 118.8 ± 29.0ab 115.1 ± 26.6ab 157.0 ± 8.0b 33.36 ± 4.87b 32.27 ± 1.07a 28.6 ± 3.1a 195.7 ± 2.9a d28F6 4.65 ± 0.06c 1.50 ± 0.10ab 22.70 ± 1.04a 63.1 ± 10.4c 49.7 ± 2.2c 90.3 ± 0.8c 10.20 ± 1.92c 24.54 ± 4.15ab 4.9 ± 1.1c 25.3 ± 1.1b Values in the table represent mean ± standard error (n = 3). Different lowercase letters indicate significant differences among treatments within the same land use type and moisture content (P < 0.05). Treatment names: d14 denotes day 14 of cultivation, d28 denotes day 28; M indicates addition of mealworm castings, 6 denotes 60% field capacity moisture content, 8 denotes 80% field capacity moisture content; D denotes croplands, L denotes grassland, F denotes forest land. Among replicates, the coefficient of variation (CV; standard deviation/mean) was calculated. When the CV exceeded 0.3, one outlier was removed from the three replicate measurements to minimize the CV as much as possible. 3.2 Analysis of the Effects of Tenebrio molitor Castings on Soil Microbial Community Diversity 3.2.1 Alpha Analysis of Soil Microbial Communities Analysis of diversity indices for the different treatments is shown in Table 4 . When insect frass was added to different land-use types, the bacterial Shannon index decreased significantly in the cultivated land (DL6) and grassland (LN6) treatments compared with the blank control, with reductions ranging from 4.67% to 7.83%. Whereas the Shannon index for fungi showed a significant increase in the forest land (FT6) treatment, with the rate of increase intensifying over time, rising from 1.75% on day 14 to 5.98% on day 28. The variation in the Shannon index for fungi across different treatments was similar to that observed for bacteria; the decline in the DL treatments was relatively small, ranging from 12.1% to 23.6%; the LN treatment showed a larger decrease, at 30.6% on day 14 and 25.2% on day 28; the FT treatment showed a slight increase after the addition of insect faeces, but this was not statistically significant. Regarding the bacterial Simpson’s index, following the addition of insect faeces, only the cultivated land treatment showed a significant decrease compared with the blank control; the grassland treatment showed no significant change, whilst the woodland treatment showed a significant increase on day 28. The Simpson index for fungi showed more pronounced changes. Following the addition of insect faeces, the arable land treatment showed no significant change; the grassland treatment exhibited a significant decrease of 13.0% on day 14, which improved slightly by day 28 compared to day 14, yet the treatment with added insect faeces remained significantly lower than the blank control, with a decrease of 7.42%; The trend in the forest plot was the opposite to that of the other treatments; the fungal Simpson’s index in the insect faeces-added treatment was significantly higher than that of the blank control, with an increase of 33.4%–44.3%. The changes in the bacterial ACE and Chao-1 indices were consistent: following the addition of insect faeces, the arable land treatment showed a significant decrease compared to the blank control; the grassland treatment also showed a significant decrease compared to the blank control; however, there was no significant difference among the forest plot treatments. The trends in fungal ACE and Chao-1 indices were consistent: following the addition of insect frass, the arable land, forest land and grassland treatments all showed a significant decrease compared to the control, with a reduction ranging from 21.6% to 57.3%. Translated In cultivated land, the effects of different moisture content treatments on microbial diversity also varied considerably. In the treatment with a moisture content of 80% of field capacity (DL8), the treatment supplemented with insect frass showed, on day 14, that the bacterial Shannon, bacterial Simpson, bacterial ACE, bacterial Chao1, the fungal ACE and Chao-1 indices were significantly lower than those of the control treatment; however, by day 28, only the fungal ACE and Chao-1 indices were significantly lower than those of the control treatment; whereas there were no significant differences between treatments in the fungal Simpson and Shannon indices. Table 4 Assessment of Soil Microbial Diversity Under Different Treatments Treatments Bacterial Shannon Index Bacterial Simpson Index Bacterial Chao 1 Index Bacterial ACE Index Fungal Shannon Index Fungal Simpson Index Fungal Chao 1 Index Fungal ACE Index d0DL 6.81 ± 0.06a 1.00 ± 0.00a 2256 ± 35a 2256 ± 35a 3.27 ± 0.09a 0.90 ± 0.01a 318 ± 24ab 318 ± 24ab d0LN 7.20 ± 0.08a 1.00 ± 0.00a 2583 ± 142a 2583 ± 142a 4.38 ± 0.08a 0.96 ± 0a 515 ± 22a 515 ± 22a d0FT 6.63 ± 0.11a 1.00 ± 0.00ab 1872 ± 403a 1872 ± 403a 2.36 ± 0.22a 0.70 ± 0.05a 286 ± 27a 286 ± 27a d14DL60M 6.10 ± 0.1c 0.99 ± 0.00cd 1472 ± 105c 1472 ± 105c 2.51 ± 0.2b 0.82 ± 0.03a 191 ± 19c 191 ± 19c d14DL60 6.40 ± 0.04b 1.00 ± 0.00b 1683 ± 83b 1683 ± 83b 2.98 ± 0.1ab 0.86 ± 0.02a 262 ± 23ab 262 ± 23ab d14DL80M 6.08 ± 0.34c 0.99 ± 0.00b 1300 ± 276c 1300 ± 276c 2.56 ± 0.16c 0.83 ± 0.04ab 177 ± 20c 177 ± 20c d14DL80 6.60 ± 0.11ab 1.00 ± 0.00a 1772 ± 120b 1772 ± 120b 2.92 ± 0.13abc 0.83 ± 0.04ab 281 ± 48ab 281 ± 48ab d14LN60M 6.04 ± 0.26d 1.00 ± 0.00d 1602 ± 212d 1602 ± 212d 2.68 ± 0.05d 0.82 ± 0.01c 172 ± 38d 172 ± 38d d14LN60 6.51 ± 0.05bc 0.99 ± 0.00cd 2036 ± 130bc 2036 ± 130bc 3.87 ± 0.05b 0.95 ± 0a 324 ± 75c 324 ± 75c d14FT60M 6.29 ± 0.04ab 1.00 ± 0.00ab 1390 ± 54b 1390 ± 54b 1.42 ± 0.17b 0.58 ± 0.05ab 115 ± 7c 115 ± 7c d14FT60 6.18 ± 0.34b 0.99 ± 0.00b 1345 ± 270b 1345 ± 270b 1.3 ± 0.28b 0.44 ± 0.11b 174 ± 27b 174 ± 27b d28DL60M 6.04 ± 0.06c 0.99 ± 0.00d 1471 ± 75c 1471 ± 75c 2.64 ± 0.59b 0.82 ± 0.11a 225 ± 42bc 225 ± 42bc d28DL60 6.38 ± 0.1b 0.99 ± 0.00bc 1688 ± 88b 1688 ± 88b 3.45 ± 0.13a 0.91 ± 0.02a 287 ± 43a 287 ± 43a d28DL80M 6.39 ± 0.07bc 1.00 ± 0.00ab 1505 ± 87bc 1505 ± 87bc 2.65 ± 0.43bc 0.81 ± 0.07b 245 ± 54bc 245 ± 54bc d28DL80 6.45 ± 0.16b 1.00 ± 0.00a 1616 ± 123b 1616 ± 123b 3.02 ± 0.09ab 0.83 ± 0.02ab 322 ± 50a 322 ± 50a d28LN60M 6.31 ± 0.13c 0.99 ± 0.00bc 1765 ± 128cd 1765 ± 128cd 3.31 ± 0.42c 0.90 ± 0.05b 181 ± 4d 181 ± 4d d28LN60 6.75 ± 0.04b 1.00 ± 0.00b 2176 ± 128b 2176 ± 128b 4.42 ± 0.11a 0.97 ± 0.01a 424 ± 4b 424 ± 4b d28FT60M 6.60 ± 0.17a 1.00 ± 0.00a 1772 ± 265ab 1772 ± 265ab 1.72 ± 0.16b 0.65 ± 0.05a 191 ± 13b 191 ± 13b d28FT60 6.23 ± 0.17b 0.99 ± 0.00ab 1446 ± 110ab 1446 ± 110ab 1.49 ± 0.39b 0.45 ± 0.12b 273 ± 26a 273 ± 26a Values in the table represent mean ± standard error (n = 3). Different lowercase letters indicate significant differences among treatments within the same land use type and moisture content (P < 0.05). Treatment designations: d14 denotes day 14 of cultivation, d28 denotes day 28; M indicates addition of mealworm castings, 6 denotes 60% field capacity moisture content, 8 denotes 80% field capacity moisture content; D denotes croplands, L denotes grassland, F denotes forest land. 3.2.2 Beta Analysis of Soil Microbial Communities The RDA analysis of bacterial communities is shown in Fig. 2 (a). RDA1 explained 46.34% of the variation, RDA2 explained 32.17%, and the cumulative explanation reached 78.51%. The physicochemical factors pH, available phosphorus (AvbP), and available potassium (AvbK) had the greatest effects on the bacterial community. Overall, the bacterial community composition in soils amended with insect frass was clearly differentiated from that in soils without insect frass addition. In the grassland (LN) and forest land (FT) treatments, the bacterial communities in the insect frass-amended treatments were clearly differentiated from those in the blank controls; the differences between treatments were more pronounced on day 14 and became smaller on day 28. In the cropland treatment (DL), the treatment with 60% field water-holding capacity (DL 6) showed clear differentiation from the blank control after insect frass addition, but the degree of differentiation was smaller than that in the grassland and forest land treatments. Compared with the blank control, the treatment with 80% field water-holding capacity (DL 8) showed relatively clear differentiation on day 14 after insect frass addition, whereas the differentiation was not obvious on day 28. Among the three different land-use types, the forest land treatment showed the most pronounced difference from the blank control after insect frass addition. RDA analysis of fungal communities is shown in Fig. 2 (b). RDA1 explained 85.68% of the variation, RDA2 explained 9.41%, and the cumulative explanation reached 95.09%; the explanatory power of the RDA1 axis was much greater than that of the RDA2 axis. Among the physicochemical factors, pH had the greatest effect on the fungal community, followed by available phosphorus and available potassium. Compared with the blank control, all treatments amended with insect frass showed clear differentiation along the RDA2 axis, indicating obvious differences in fungal communities. In the grassland treatments, compared with the blank control, the differences between the insect frass-amended treatments were more pronounced on day 14 and became smaller on day 28. In the forest land treatments, compared with the blank control, the differences in fungal communities in the insect frass-amended treatments became increasingly greater over time. In the cropland treatments, for the treatment with 60% field water-holding capacity, the difference from the blank control did not decrease obviously over time after insect frass addition. Compared with the DL 6 treatment, the treatment with 80% field water-holding capacity showed relatively small differences in fungal communities after insect frass addition. In addition to pH, available phosphorus, and available potassium, total nitrogen, total carbon, and soil soluble carbon also had relatively large effects on fungal community composition. 3.3 Effects of Tenebrio molitor Castings on Soil Microbial Community Structure and Composition In this study, we performed ASV (Amplicon Sequence Variant) clustering analysis on the valid sequences from all samples. Following species annotation, the top 10 bacterial phyla were selected for analysis, as shown in Fig. 3 (a). The results showed that the bacterial communities exhibited clear differences at the phylum level under different soil types and incubation treatments, and that incubation time, moisture content, and insect frass addition had different effects on the relative abundances of the dominant bacterial phyla. Overall, the dominant bacterial phyla in samples from all treatments mainly included Pseudomonadota, Actinomycetota, Acidobacteriota, Bacteroidota, Bacillota, and Gemmatimonadota, but the variation trends were not consistent among different soils. In the cropland treatments, the insect frass-amended treatment at 60% field water-holding capacity (DL 6) showed high abundances of Pseudomonadota and Bacteroidota on days 14 and 28. Over time from day 14 to day 28, the abundance of Gemmatimonadota further increased under insect frass addition. In the high-moisture treatment (DL 8) amended with insect frass, the abundances of Pseudomonadota, Bacteroidota, and Gemmatimonadota were high on days 14 and 28. However, on day 28, the abundance of Pseudomonadota decreased, whereas that of Bacillota increased under insect frass addition. Over time from day 14 to day 28, Pseudomonadota showed a decreasing trend, whereas Gemmatimonadota showed an increasing trend under insect frass addition. In the grassland (LN) and forest land (FT) treatments, the 60% field water-holding capacity treatment amended with insect frass showed that Pseudomonadota and Acidobacteriota maintained high abundances on days 14 and 28. Over time from day 14 to day 28, the abundance of Acidobacteriota further increased under insect frass addition. Analysis of the top 10 fungal phyla at the phylum level is shown in Fig. 3 (b). In the cropland treatments, the insect frass-amended treatment at 60% field water-holding capacity (DL 6) showed a relatively high abundance of Ascomycota on day 14. On day 28, Ascomycota still maintained a high abundance under insect frass addition, whereas Basidiomycota and Others were relatively higher in the control treatment. Over time from day 14 to day 28, the major fungal phyla showed little overall change under insect frass addition. In the high-moisture treatment (DL 8) amended with insect frass, Ascomycota showed a high abundance on day 14, whereas Others showed a relatively low abundance. On day 28, Ascomycota still remained the absolutely dominant phylum under insect frass addition, while Basidiomycota and Mortierellomycota increased slightly. Over time from day 14 to day 28, Basidiomycota and Others increased slightly under insect frass addition. In the grassland treatment (LN), the 60% field water-holding capacity treatment amended with insect frass showed a high abundance of Ascomycota on day 14, whereas Basidiomycota and Others were relatively higher in the control treatment. On day 28, Ascomycota still maintained a high abundance under insect frass addition. Over time from day 14 to day 28, Ascomycota decreased slightly, whereas Basidiomycota and Others increased slightly under insect frass addition. In the forest land treatment (FT), the 60% field water-holding capacity treatment amended with insect frass showed that the abundance of Ascomycota was higher than that in the control on day 14, whereas the abundance of Basidiomycota was lower than that in the control. On day 28, this change became more obvious: under insect frass addition, Ascomycota further increased, whereas Basidiomycota further decreased; in the control treatment, Basidiomycota consistently maintained a high abundance. Over time from day 14 to day 28, Ascomycota showed an increasing trend, whereas Basidiomycota showed a decreasing trend under insect frass addition. The composition and relative abundance of bacterial communities at the genus level under different treatments are shown in Fig. 4 . In the cropland treatment, under 60% field water-holding capacity (DL 6), the insect frass-amended treatment showed higher abundances of Chryseolinea, Steroidobacter, Pontibacter, Ohtaekwangia, Lysobacter, Nonomuraea, and Chitinophaga than the blank control. From day 14 to day 28, the abundances of Chryseolinea and Steroidobacter further increased in the insect frass-amended treatment, whereas the abundance of Pontibacter decreased slightly, with no significant changes in the blank control. Under 80% field water-holding capacity (DL 8), the insect frass-amended treatment showed increased abundances of Ohtaekwangia and Lysobacter, whereas Ligilactobacillus and Romboutsia decreased; over time, Pontibacter and Ohtaekwangia showed a decreasing trend in the insect frass-amended treatment. In the grassland treatment (LN), under 60% field water-holding capacity, the insect frass-amended treatment showed higher abundances of Lysobacter, Nonomuraea, Chitinophaga, and Steroidobacter than the blank control, whereas Micromonospora and Ligilactobacillus decreased; with prolonged incubation, the abundances of Lysobacter and Micromonospora decreased markedly, whereas those of Nonomuraea and Chitinophaga increased明显. In the forest land treatment (FT), under 60% field water-holding capacity, the insect frass-amended treatment showed markedly higher abundances of Chitinophaga, Sphingomonas, and Burkholderia-Caballeronia-Paraburkholderia than the blank control, whereas Acidothermus, Candidatus Koribacter, Bryobacter, and Candidatus Solibacter decreased. Over time, the abundances of Burkholderia-Caballeronia-Paraburkholderia and Massilia decreased markedly in the insect frass-amended treatment, whereas the blank control showed little change. The composition and relative abundance of fungal communities at the genus level under different treatments are shown in Fig. 5 . In the cropland treatment, under 60% field water-holding capacity (DL 6), the insect frass-amended treatment showed increased abundances of Mortierella and Chaetomiaceae_gen_Incertae_sedis, whereas the abundance of Stachybotrys decreased. From day 14 to day 28, the abundance of Mortierella continued to increase in the insect frass-amended treatment, whereas Stachybotrys in the control treatment increased slightly. In the high-moisture treatment (DL 8), the insect frass-amended treatment showed an increased abundance of Aspergillus, whereas the abundances of Stachybotrys and Chaetomium decreased; over time, the abundances of Zopfiella and Pezizales_gen_Incertae_sedis increased in the insect frass-amended treatment. In the grassland treatment (LN), under 60% field water-holding capacity, insect frass addition markedly increased the abundances of Striaticonidium, Pseudorhypophila, and Humicola, whereas Scedosporium, Gliocephalotrichum, and Oliveonia decreased. With prolonged incubation, Striaticonidium and Humicola decreased slightly, whereas Periconia and Exserohilum increased in the insect frass-amended treatment; in the control treatment, Trichoderma, Fusarium, Scedosporium, and Gliocephalotrichum showed increasing trends, whereas Exserohilum, Neopestalotiopsis, Serendipitaceae_gen_Incertae_sedis, and Ovatospora decreased. In the forest land treatment (FT), under 60% field water-holding capacity, insect frass addition increased the abundance of Penicillium, whereas Trichoderma, Mariannaea, and Saitozyma decreased. Over time, on day 28 compared with day 14, Parachaetomium further increased in the insect frass-amended treatment, whereas the abundances of Trichoderma and Mariannaea decreased in the control treatment. 3.4 Correlation Analysis Between Soil Microbial Communities and Physicochemical Factors The correlations among physicochemical indicators and different variable factors, as well as their correlations with bacterial community abundance changes, are shown in Fig. 6 . The addition of insect frass generally increased the abundances of three bacterial phyla, Pseudomonadota, Bacteroidota, and Chloroflexota, in soil; high soil moisture promoted an increase in the abundance of Chloroflexota, and the abundance of Chloroflexota in soil also increased over time. In all treatments, total soluble carbon (TSC) and total soluble organic carbon (TSOC) were significantly positively correlated with the abundance of Actinomycetota; soil ammonium nitrogen (Ammonium) was significantly positively correlated with the abundance of Pseudomonadota; soil nitrate nitrogen (Nitrate) was positively correlated with the abundance of Thermoproteota; soil available potassium (AvbK) was significantly positively correlated with the abundances of Acidobacteriota and Actinomycetota; soil alkali-hydrolyzable nitrogen (AvbN) was significantly positively correlated with the abundances of Pseudomonadota and Actinomycetota; soil available phosphorus (AvbP) was significantly positively correlated with the abundances of Bacteroidota and Gemmatimonadota; soil total nitrogen (TN) was significantly positively correlated with the abundances of Actinomycetota and Bacteroidota; soil total carbon (TC) was significantly positively correlated with the abundances of Actinomycetota and Chloroflexota; and soil pH was significantly positively correlated with the abundances of Acidobacteriota, Gemmatimonadota, Candidatus Eremiobacterota, and Verrucomicrobiota. The correlations among physicochemical indicators and different variable factors, as well as their correlations with fungal community abundance changes, are shown in Fig. 7 . The addition of insect frass generally significantly increased the abundance of Fungi_phy_Incertae_sedis in soil; high soil moisture promoted increases in the abundances of Mucoromycota and Fungi_phy_Incertae_sedis, and the abundance of Fungi_phy_Incertae_sedis in soil also increased over time. In all treatments, total soluble carbon (TSC) was significantly positively correlated with the abundance of Glomeromycota; total soluble organic carbon (TSOC) was significantly positively correlated with the abundance of Basidiomycota; soil nitrate nitrogen (Nitrate) was positively correlated with the abundances of Mortierellomycota and Zoopagomycota; soil available potassium (AvbK) was significantly positively correlated with the abundances of Ascomycota, Mortierellomycota, and Mucoromycota; soil alkali-hydrolyzable nitrogen (AvbN) was significantly positively correlated with the abundance of Basidiomycota; soil available phosphorus (AvbP) was significantly positively correlated with the abundances of Ascomycota, Basidiomycota, and Mortierellomycota; soil total nitrogen (TN) and soil total carbon (TC) were significantly positively correlated with the abundance of Glomeromycota; and soil pH was significantly positively correlated with the abundances of Ascomycota, Basidiomycota, and Mucoromycota. 3.5 Overall effects on soil after applying mealworm castings Figure 8 shows the PLS-SEM results for the effects of insect frass application on soil properties across the three land-use types under the condition that soil moisture was maintained at 60% of field water-holding capacity. The addition of insect frass reduced total nitrogen and total carbon contents and soluble soil nutrients (soluble carbon and nitrogen) to a certain extent, significantly increased available soil nutrients (available nitrogen, phosphorus, and potassium), and gradually increased soil pH. After the application of mealworm frass, the dominant bacterial phyla in the soil mainly belonged to the functional groups involved in carbon cycling and in nitrogen and phosphorus cycling. Total nutrients, soluble nutrients, and available nutrients were all positively correlated with both microbial functional groups. Among them, total nutrients showed the strongest significant positive correlation with the bacterial communities involved in carbon cycling (“Actinomycetota”, “Bacteroidota”, “Gemmatimonadota”), while soluble nutrients showed the strongest significant positive correlation with the carbon-cycling bacteria (“Gemmatimonadota”, “Pseudomonadota”). 3.6 The Effect of Mealworm Castings on Soil as Determined by Soil Mechanical Composition By calculating the rate of change in various indicators between the treatment applying mealworm castings and its blank control, SEM was constructed. Soil texture effects on the performance of yellow mealworm castings are shown in Fig. 9 . Total soil nitrogen and total soil carbon were positively correlated only with sand content, and the correlations were weak. Sand content was significantly positively correlated with soil available nutrients and soil pH (p < 0.05), and negatively correlated with total soluble carbon, although the correlation with total soluble carbon was weak. Silt content was significantly positively correlated with soil pH (p < 0.05) and negatively correlated with soil available nutrients (p < 0.05) and total soluble carbon. Clay content was significantly negatively correlated with soil pH (p < 0.05) and positively correlated with total soluble carbon. 4 Discussion 4.1 Effects of Different Treatments on Soil Physicochemical Properties After mealworm castings were added, changes in soil physicochemical properties differed among land-use types. This variation was partly attributable to differences in baseline soil properties, which caused mealworm castings to affect different soils to different extents. Following the addition of mealworm castings, soil pH generally shifted toward neutrality (approximately 7.0). As an organic amendment, mealworm castings can buffer soil pH and, to some extent, increase pH in acidic soils (Li, D. R. et al ., 2013; Li, Q. J. et al ., 2014). Incorporation of mealworm castings can influence soil pH and nutrient status through multiple pathways. Specifically, mealworm castings can increase soil pH (particularly in acidic soils), enhance soil organic carbon and total nitrogen, and increase available phosphorus and available potassium, while also improving soil structure and microbial activity (Bai, Z. et al., 2018 ; Antoniadis, V. et al., 2023 ). In cropland treatments, changes in soil soluble carbon and soluble organic carbon after mealworm casting application showed distinct temporal patterns relative to the blank control under 80% versus 60% of field capacity. Higher moisture can partly slow microbial decomposition of organic matter (Song, D. et al., 2022 ). It also promotes the accumulation of soil ammonium nitrogen while reducing nitrate nitrogen content. As an organic soil conditioner, mealworm castings significantly enhance soil easily decomposable carbon content by day 14. Compared to day 14, easily decomposable carbon decreases by day 28, while readily available nutrients begin to increase gradually. This suggests that, during the initial phase after application, mealworm castings undergo preliminary microbial decomposition. Over time, microbial activity progressively mineralizes the castings and releases plant-available nutrients (Zunzunegui, I. et al., 2024 ; Antoniadis, V. et al., 2023 ). However, high moisture content (80% of field capacity) reduced the overall effects of mealworm castings on soil physicochemical properties and slowed microbial mineralization and nutrient release. 4.2 Effects of Different Treatments on Microbial Community Characteristics After insect feces was added, the Shannon and Simpson indices increased in forest land (FT) but decreased in croplands (DL) and grassland (LN). These contrasting responses may be driven by multiple factors, including differences in baseline soil conditions, nutrient status, microbial community structure, and niche competition among land-use types (Demenois, J. et al., 2020 ; Huang, Q. et al., 2024 ). In forest land ecosystems, baseline microbial diversity may be relatively low, potentially due to environmental constraints specific to forest ecosystems. When mealworm castings are applied, they act as an organic amendment and supply nutrients such as organic matter, nitrogen, and phosphorus (Li, N. et al., 2024 ; Gurmessa, B. et al., 2021 ). These nutrient inputs may stimulate the growth of microbial taxa that were previously inactive or rare in forest soils, thereby increasing species richness and evenness and ultimately elevating Shannon and Simpson indices (Demenois, J. et al., 2020 ; Li, N. et al., 2024 ). In croplands and grasslands, where baseline diversity is relatively high, nutrients in mealworm castings may preferentially promote the proliferation of already dominant taxa. Moreover, the combined effects of amendment inputs, disturbance to the original soil environment, and potential nutrient surpluses may intensify competitive exclusion, leading to reduced community evenness and, consequently, decreases in the Shannon and Simpson indices (Liu, Z. et al., 2022 ; Wang, X. et al., 2023 ; Zhang, M. et al., 2024 ). Changes in soil physicochemical properties can strongly shape bacterial community composition. After mealworm feces addition, soil pH, available phosphorus, readily available potassium, and soluble carbon and nitrogen exerted the strongest effects on bacterial communities. Compared with the blank controls, bacterial communities in grassland and forest soils diverged more markedly by day 14, indicating a stronger early-phase response following amendment application. In contrast, bacterial community shifts in cropland soils were less pronounced than those in grassland and forest soils. This difference may be attributable to the relatively higher baseline nutrient status of cropland soils, such that amendment-induced changes in physicochemical properties exert a smaller incremental effect on microbial communities than in forest soils. For example, studies in temperate grassland ecosystems have shown that soil bacterial community composition can be more strongly influenced by nutrient availability than by pH (Zhang, H. et al., 2024 ; Tian, H. et al., 2017 ). Compared to the control, fungal communities in both forest and grassland soils under the insect dung treatment were more strongly associated with soil available phosphorus, available potassium, and pH on day 14. However, by day 28, the primary factors driving differentiation between the two treatments shifted to soluble carbon, soluble organic carbon, total carbon, ammonium nitrogen and alkaline-hydrolyzable nitrogen. This shift may be explained by temporal changes in substrate availability following amendment addition. Immediately after application, mealworm castings supply abundant nutrients that favor rapid microbial growth. Over time, the introduced carbon is transformed into more labile fractions and undergoes continued mineralization. Consequently, fungal taxa that initially dominate the decomposition of readily available substrates may decline in relative dominance. The functional emphasis of the fungal community may therefore shift from exploiting readily available nutrients toward utilizing more complex carbon sources and/or adapting to conditions with a lower carbon-to-nitrogen ratio (Mao, X. et al., 2024 ). 4.3 Effects of Different Treatments on Microbial Community Composition The addition of insect feces overall increased the abundance of Pseudomonadota, Actinomycetota, Acidobacteriota, Bacteroidota, Bacillota and Gemmatimonadota within the bacterial community. Mealworm frass is rich in incompletely digested plant polysaccharides, chitin, short-chain fatty acids, and microbial metabolites, which provide soil bacteria with diverse carbon and nitrogen sources (Verardi, A. et al., 2025 ; Nurfikari, A. et al., 2024 ). For example, Bacteroidota and Pseudomonadota , which are capable of degrading complex polysaccharides, as well as Actinomycetota , which can utilize recalcitrant organic matter, may gain a competitive advantage because of the presence of these substrates, thereby increasing in abundance (Nurfikari, A. et al., 2024 ). The addition of mealworm frass alters the physicochemical properties of soil, such as pH, moisture content, redox potential, and oxygen diffusion rate (Nurfikari, A. et al., 2024 ; Nogalska, A. et al., 2024 ). These changes in the microenvironment have selective effects on the growth of different bacterial groups. Mealworm frass may contain bioactive substances produced by the insects themselves, such as antimicrobial peptides, chitinases, and metabolites, and these substances may exert direct or indirect effects on the soil microbial community, for example, by inhibiting certain pathogens or promoting the growth of beneficial bacteria (Rizou, E. et al., 2025 ). The addition of mealworm frass increases the abundance of Ascomycota in soil. Mealworm frass contains chitin derived from mealworm molting, which is a nitrogen-containing polysaccharide (Verardi, A. et al., 2025 ; Blakstad, J. I. et al ., 2023). Many groups within Ascomycota , such as Fusarium and Trichoderma , are known for their strong chitin-degrading ability (Zhang, J. et al., 2024 ). The ability to utilize chitin efficiently may confer a competitive advantage on these ascomycetous fungi in environments amended with mealworm frass, thereby increasing their relative abundance. Overall, the addition of mealworm frass increased the abundance of Chitinophaga in soil. This phenomenon was mainly attributed to the high chitin content in mealworm frass, which provided sufficient carbon and nitrogen sources for the growth of Chitinophaga (Verardi, A. et al., 2025 ; Zunzunegui, I. et al., 2024 ). In the cropland treatment, insect frass addition promoted increases in the abundances of bacterial communities such as Chryseolinea , Steroidobacter , Pontibacter , Ohtaekwangia , Lysobacter , Nonomuraea , and Chitinophaga . The increases in these bacterial groups suggest that the soil community was shifting toward enhanced nutrient transformation, biological control, and organic matter mineralization, thereby improving overall soil productivity (Nie, J. et al., 2025 ; Nurfikari, A. et al., 2024 ). In the grassland treatment, insect frass application led to increased abundances of the three saprophytic filamentous fungi Striaticonidium , Pseudorhypophila , and Humicola . These fungi play important roles in the decomposition of recalcitrant organic matter (such as lignin and keratin) and in humification, thereby contributing to the accumulation and stabilization of soil organic carbon (Jeffery, S. et al., 2022 ; Hannula, S. E. et al ., 2021). At the same time, the abundances of genera such as Scedosporium , Gliocephalotrichum , and Oliveonia decreased, some of which are considered opportunistic pathogens or weak plant pathogens, suggesting enhanced niche competition and the formation of a disease-suppressive environment (Zhou, W. et al ., 2023; Zhao, W. et al ., 2023). In the cropland and grassland treatments, the addition of mealworm frass significantly increased the abundances of Striaticonidium , Pseudorhypophila , and Humicola , whereas the abundances of Scedosporium , Gliocephalotrichum , and Oliveonia decreased. In the forest land treatment, the addition of mealworm frass increased the abundance of Penicillium , whereas the abundances of Trichoderma , Mariannaea , and Saitozyma decreased. Increases in fungi such as Striaticonidium and Humicola indicate that the degradation and mineralization of soil organic matter were enhanced, which helps accelerate nutrient cycling and convert unstable organic carbon into stable humus, thereby improving soil fertility and carbon sequestration capacity (Yan, H. et al ., 2023; Miao, Y. et al ., 2022). Extracellular enzymes and microbial products secreted by fungi such as Humicola help form and stabilize soil aggregates, improve soil physical structure, enhance soil water- and nutrient-holding capacity, and increase resistance to erosion (Readyhough, T. et al ., 2021; Schroeder, J. et al ., 2021). Increases in beneficial fungi such as Pseudorhypophila and Penicillium, together with decreases in potentially harmful fungi such as Scedosporium, Gliocephalotrichum, and Oliveonia, may enhance soil resistance to pathogens and reduce the risk of plant disease occurrence (Yan, W. et al., 2024 ; Tao, C. et al ., 2023). These changes indicate that mealworm frass, as an organic amendment, can effectively regulate soil microbial communities, particularly fungal communities, thereby optimizing soil ecological functions and supporting sustainable agricultural practices. 4.4 Effects of Applying Mealworm Castings on Soil Through iterative modeling, we found that the application of mealworm castings introduced abundant nutrients and bioactive substances into the soil, thereby influencing the abundance and activity of the soil microbial community. Microorganisms promoted soil nutrient cycling through their metabolic activities. Following application, microorganisms associated with carbon, nitrogen, and phosphorus cycling became dominant in the soil. These microbes mineralized large organic molecules, converting them into soluble carbon and nitrogen and increasing available phosphorus. Previous studies indicate that mealworm castings effectively elevate available phosphorus levels in soil. Therefore, applying mealworm castings enhances soil nutrient cycling capacity, converting nutrients difficult for plants to utilize into forms that can be readily absorbed by plants (Watson, C. et al., 2021 ; Rumbos, C. I. et al ., 2025; Verardi, A. et al., 2025 ). It can also gradually improve soil pH. 4.5 Effects of Soil Moisture and Mechanical Composition on the Performance of Yellow Mealworm Castings By calculating the relative change rates between the mealworm casting treatment and the blank control and applying iterative modeling, we found that excessively high soil moisture accelerated the increase in soil ammonium nitrogen following mealworm casting application. At the same time, high moisture inhibited the mineralization of mealworm castings by day 14. Only by day 28 did the castings begin to mineralize gradually, at which point soil available nutrients started to increase; however, the increase in soil nitrate nitrogen remained suppressed. Excessive moisture likely promotes ammonification of organic nitrogen by reducing oxygen availability, leading to ammonium accumulation. It may also inhibit aerobic mineralization of organic matter, thereby delaying decomposition. Finally, high moisture strongly suppresses nitrification, limiting nitrate formation (Keiluweit, M. et al., 2018 ; Sun, S. et al., 2020 ; Sriraj, P. et al., 2022 ). Over time, microbial communities may adapt and/or initiate partial anaerobic mineralization, which could restore overall decomposition to some extent; nevertheless, nitrate accumulation is likely to remain constrained under persistently high moisture conditions. In soils with different mechanical composition, higher sand contents tend to promote the mineralization of mealworm castings into plant-available nutrients. By contrast, higher silt content was unfavorable for the accumulation of soluble nutrient pools (e.g., total soluble carbon and total soluble nitrogen) and available nutrients. Excessively high clay contents can inhibit both the mineralization of mealworm castings and their pH-modifying effects. Overall, soil texture regulates the mineralization rate of mealworm castings and their effects on soil nutrients and pH by shaping porosity, aeration, water-holding capacity, microbial accessibility to substrates, and pH buffering capacity. Soils with higher sand contents typically have greater porosity and better aeration, which enhance oxygen diffusion and support aerobic microbial activity. In sandy soils, aerobic microorganisms can more efficiently decompose organic matter in mealworm castings and mineralize it into plant-available nutrients such as alkaline-hydrolyzable nitrogen and readily available phosphorus (Ahmed, W. et al., 2024 ; Grzyb, A. et al., 2020 ). Soils with higher silt contents, owing to their relatively large specific surface area, often exhibit stronger adsorption capacity and provide attachment sites for microorganisms, thereby promoting the mineralization of total nitrogen and total carbon in soil (Mao, H.-R. et al ., 2024). In contrast, soils with excessively high clay contents may suppress microbial activity because of poorer aeration and higher water-holding capacity, thereby reducing the decomposition of mealworm castings (Ahmed, W. et al., 2024 ). Moreover, negative charges on clay minerals and organic matter surfaces can adsorb cations, influencing soil buffering capacity and pH dynamics (Jeon, I. et al., 2023 ). In soils with high clay content, the inherently stronger buffering capacity may make pH changes induced by mealworm castings less pronounced. For soils of different textures, their specific characteristics should be considered to optimize the application strategy of mealworm castings, maximizing its ecological benefits as an organic fertilizer source. 5 Conclusion Overall, applying mealworm castings increased various available nutrients in the soil. However, across different land use types, baseline soil nutrient levels affected the effectiveness of mealworm castings. The application of mealworm castings involved relatively low human intervention and showed the most pronounced improvements in relatively nutrient-poor soils, while also producing beneficial effects in nutrient-rich soils. After insect dung application, the abundance of soil saprotrophic microorganisms and other beneficial microbial communities increased to some extent, whereas the abundance of pathogenic microorganisms decreased, which substantially enhanced soil health and improved crop disease resistance. The recommended moisture condition for applying mealworm castings was 60% of field capacity, because excessively high moisture slowed mineralization and decomposition of Tenebrio molitor castings and delayed the time required for them to become effective in soil. Higher sand and silt contents facilitated the mineralization and decomposition of Tenebrio molitor castings, whereas high clay content inhibited the effectiveness of mealworm castings. This study elucidated how mealworm castings influenced soil physicochemical properties and microbial communities across soils differing in land-use type, texture, and moisture regime. However, the present work focused on soil responses and did not evaluate outcomes during crop cultivation. Evidence remains limited regarding optimal application rates required to maximize overall benefits under different land-use types and moisture conditions. In addition, the potential role of leaching was not considered when assessing texture-dependent effects. Future studies should address these gaps through complementary experiments to provide more robust and comprehensive guidance for the agricultural use of mealworm castings. Declarations Author Responsibility Statement We, the authors of this paper, hereby solemnly declare once more that we confirm this research constitutes original work, with all experimental data being first-hand data obtained through standardised experiments. This study has not involved any form of academic misconduct, including but not limited to data falsification, plagiarism, multiple submissions, or duplicate publication. All references have been properly cited. Conflict of Interest Statement We declare that there are no competing financial interests or personal relationships relevant to this research. Author Contribution Z:Responsible for experimental design, sample collection and mechanical composition analysis; completed moisture infiltration rate measurements and data collation for the earthworm compost treatment group; drafted the initial manuscript. Z:Participated in determining soil physical and chemical properties, sample collection, assisted in collating data across different land use types, conducted data plotting, chart visualisation, and paper revisions. L:Responsible for earthworm biomass monitoring and soil microbial community analysis, collated comparative data across different land use types and performed data processing. W and L:Assisted with earthworm biomass monitoring and preliminary microbial community analysis. J and J:Assisted with soil sample collection and data organisation. Z:As supervising tutor, provided research direction, land use typedata support, and field sampling technical guidance. X:As corresponding author, guided research direction, provided conceptual and technical support, and reviewed the manuscript. Acknowledgements This work was supported by the Natural Science Foundation of Guangdong Province (2023A1515010593). Data Availability All data generated or analyzed during this study are included in this published article and its supplementary information files References He L, Zhang Y, Ding M et al (2021) Sustainable strategy for lignocellulosic crop wastes reduction by Tenebrio molitor linnaeus (mealworm) and potential use of mealworm frass as a fertilizer[J]. J Clean Prod, 325 Zajicova K, Chuman T (2019) Effect of land use on soil chemical properties after 190 years of forest to agricultural land conversion[J]. Soil Water Res 14(3):121–131 Adeniji A, Huang J, Li S, Lu X, Guo R (2024) Hot viewpoint on how soil texture, soil nutrient availability, and root exudates interact to shape microbial dynamics and plant health. 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Also discoverable on Platform About Our Team In Review Editorial Policies Advisory Board Help Center Resources Author Services Accessibility API Access RSS feed Manage Cookie Preferences © Research Square 2026 | ISSN 2693-5015 (online) Privacy Policy Terms of Service Do Not Sell My Personal Information {"props":{"pageProps":{"initialData":{"identity":"rs-8998912","acceptedTermsAndConditions":true,"allowDirectSubmit":true,"archivedVersions":[],"articleType":"Research Article","associatedPublications":[],"authors":[{"id":599823568,"identity":"4d96debe-11dc-46f3-9cca-1cd5c74f53f5","order_by":0,"name":"Zhi-Shen Hong","email":"","orcid":"","institution":"South China Agricultural University","correspondingAuthor":false,"prefix":"","firstName":"Zhi-Shen","middleName":"","lastName":"Hong","suffix":""},{"id":599823569,"identity":"a5f39aae-323f-4e49-8eaa-9752416394a6","order_by":1,"name":"Cai-Yi Zhu","email":"","orcid":"","institution":"South China Agricultural University","correspondingAuthor":false,"prefix":"","firstName":"Cai-Yi","middleName":"","lastName":"Zhu","suffix":""},{"id":599823571,"identity":"e27ef39a-7eda-416a-8610-06150dcc3161","order_by":2,"name":"Xiao-Qing Li","email":"","orcid":"","institution":"South China Agricultural University","correspondingAuthor":false,"prefix":"","firstName":"Xiao-Qing","middleName":"","lastName":"Li","suffix":""},{"id":599823572,"identity":"68eb59b5-d5f7-4b7d-bb48-15191f9230e5","order_by":3,"name":"Ruofei Wang","email":"","orcid":"","institution":"South China Agricultural University","correspondingAuthor":false,"prefix":"","firstName":"Ruofei","middleName":"","lastName":"Wang","suffix":""},{"id":599823577,"identity":"d2f53fc7-7c27-414c-8441-8fd06b6dec5d","order_by":4,"name":"Zeyu Li","email":"","orcid":"","institution":"South China Agricultural University","correspondingAuthor":false,"prefix":"","firstName":"Zeyu","middleName":"","lastName":"Li","suffix":""},{"id":599823582,"identity":"cb0898ca-063e-416e-94db-766f9f5371a6","order_by":5,"name":"Mengmeng Jiang","email":"","orcid":"","institution":"South China Agricultural University","correspondingAuthor":false,"prefix":"","firstName":"Mengmeng","middleName":"","lastName":"Jiang","suffix":""},{"id":599823583,"identity":"0f5b2468-68a1-4cd1-ad0d-beeef9c6c6dc","order_by":6,"name":"Huadong Jiang","email":"","orcid":"","institution":"South China Agricultural University","correspondingAuthor":false,"prefix":"","firstName":"Huadong","middleName":"","lastName":"Jiang","suffix":""},{"id":599823584,"identity":"593f1b10-40c8-48aa-8793-527d90ba6049","order_by":7,"name":"Yulong Zhang","email":"","orcid":"","institution":"South China Agricultural University","correspondingAuthor":false,"prefix":"","firstName":"Yulong","middleName":"","lastName":"Zhang","suffix":""},{"id":599823585,"identity":"9d74136b-3f04-4785-b43e-1c14762651ab","order_by":8,"name":"Hui-Juan Xu","email":"data:image/png;base64,iVBORw0KGgoAAAANSUhEUgAAAZAAAAAyAQMAAABI0h/eAAAABlBMVEX///8AAABVwtN+AAAACXBIWXMAAA7EAAAOxAGVKw4bAAAAu0lEQVRIiWNgGAWjYBACA4YDDAwfoBwJorUwziBRCwMDMw9JWswZz5hJ2/yxkzc4wHzwNg+DXR5BLZYNx9Kkc9uSDTccYEu25mFILibssAOHj0nnNhxg3HCAx0yah+FAYgNhLQfbpC3+HLDfcID/G7FagLYwsB1IBNrCRqyWY8mWvW3JyTMPsxlbzjFIJkLLjTOGN378sbPtO9788MabCjvCWhgkDkAZzGATCKoHAn7Cpo6CUTAKRsFIBwCjBT0lA8oPpgAAAABJRU5ErkJggg==","orcid":"","institution":"South China Agricultural University","correspondingAuthor":true,"prefix":"","firstName":"Hui-Juan","middleName":"","lastName":"Xu","suffix":""}],"badges":[],"createdAt":"2026-03-01 03:23:14","currentVersionCode":1,"declarations":"","doi":"10.21203/rs.3.rs-8998912/v1","doiUrl":"https://doi.org/10.21203/rs.3.rs-8998912/v1","draftVersion":[],"editorialEvents":[],"editorialNote":"","failedWorkflow":false,"files":[{"id":106403421,"identity":"bfc3f8ca-f9ed-4bb2-b21c-a39fc3cb3cef","added_by":"auto","created_at":"2026-04-08 09:14:15","extension":"png","order_by":1,"title":"Figure 1","display":"","copyAsset":false,"role":"figure","size":60960,"visible":true,"origin":"","legend":"\u003cp\u003ePCA Analysis of Soil Physicochemical Indicators.DL denotes croplands, FT denotes forest land, and LN denotes grassland. In the handle, d14 indicates the 14th day of cultivation, and d28 indicates the 28th day of cultivation. M denotes the addition of mealworm castings, 6 denotes a moisture content of 60% of field capacity, and 8 denotes a moisture content of 80% of field capacity. The primary axes in the figure represent the principal component (PC) values of physicochemical factors, while the secondary axes represent the PC values of the samples.\u003c/p\u003e","description":"","filename":"image1.png","url":"https://assets-eu.researchsquare.com/files/rs-8998912/v1/fa9965e04c2740f43a6ef088.png"},{"id":106403206,"identity":"1c2a8769-0375-4652-863a-852cb1647e3c","added_by":"auto","created_at":"2026-04-08 09:13:52","extension":"png","order_by":2,"title":"Figure 2","display":"","copyAsset":false,"role":"figure","size":112743,"visible":true,"origin":"","legend":"\u003cp\u003eMicrobial Community RDA Analysis.(a) represents bacterial community RDA analysis, (b) represents bacterial community RDA analysis, where “*” indicates P\u0026lt;0.05 and ≥0.01, “**” indicates P\u0026lt;0.01 and ≥0.005, and “***” indicates P\u0026lt;0.005. Primary coordinates denote environmental factor arrow RDA values, while secondary coordinates denote treatment-specific RDA values. DL denotes croplands, FT denotes forest land, LN denotes grassland. In the handle, d14 indicates day 14 of cultivation, d28 indicates day 28 of cultivation. M denotes addition of insect feces, 6 denotes moisture content at 60% field capacity, 8 denotes moisture content at 80% field capacity.\u003c/p\u003e","description":"","filename":"image2.png","url":"https://assets-eu.researchsquare.com/files/rs-8998912/v1/65ce545f38dd65d0453b4c3a.png"},{"id":106230659,"identity":"ea9f5ee1-a896-4019-bde5-2088bf31c3b4","added_by":"auto","created_at":"2026-04-06 12:32:49","extension":"png","order_by":3,"title":"Figure 3","display":"","copyAsset":false,"role":"figure","size":265538,"visible":true,"origin":"","legend":"\u003cp\u003eRelative Abundance of Soil Microbial Phyla Across Different Treatments.(a) shows the relative abundance of bacterial phyla in soils under different treatments, while (b) shows the relative abundance of fungal phyla. In treatment names, d14 denotes culture day 14 and d28 denotes culture day 28; M denotes the addition of mealworm castings; 6 indicates a moisture content of 60% of field capacity; 8 indicates a moisture content of 80% of field capacity; D represents croplands, L denotes grassland soil, and F indicates forest land soil.\u003c/p\u003e","description":"","filename":"image3.png","url":"https://assets-eu.researchsquare.com/files/rs-8998912/v1/729dd28a66753ddc76b62b6c.png"},{"id":106230661,"identity":"ceb6f300-ab6c-44b8-87cc-ce51ba2b5fae","added_by":"auto","created_at":"2026-04-06 12:32:49","extension":"png","order_by":4,"title":"Figure 4","display":"","copyAsset":false,"role":"figure","size":59597,"visible":true,"origin":"","legend":"\u003cp\u003eHeatmap of Relative Abundance Levels of Soil Bacterial Genera Across Different Treatments.In treatment designations, d14 denotes the 14th day of cultivation, and d28 denotes the 28th day of cultivation; M indicates the addition of mealworm castings, 6 denotes a moisture content of 60% of field capacity, and 8 denotes a moisture content of 80% of field capacity; D represents croplands, L denotes grassland soil, and F indicates forest land. The same applies below.\u003c/p\u003e","description":"","filename":"image4.png","url":"https://assets-eu.researchsquare.com/files/rs-8998912/v1/39ded475038e3e047467bc39.png"},{"id":106230665,"identity":"718de71c-2093-4c1e-a12c-ba25b131005d","added_by":"auto","created_at":"2026-04-06 12:32:49","extension":"png","order_by":5,"title":"Figure 5","display":"","copyAsset":false,"role":"figure","size":68093,"visible":true,"origin":"","legend":"\u003cp\u003eHeatmap of relative abundance at the genus level for soil fungi across different treatments\u003c/p\u003e","description":"","filename":"image5.png","url":"https://assets-eu.researchsquare.com/files/rs-8998912/v1/8ab95330bf7adff2108f3347.png"},{"id":106402986,"identity":"bc8f9994-cf21-4163-b967-02c83048e6a9","added_by":"auto","created_at":"2026-04-08 09:13:19","extension":"png","order_by":6,"title":"Figure 6","display":"","copyAsset":false,"role":"figure","size":122434,"visible":true,"origin":"","legend":"\u003cp\u003eCorrelation Analysis Between Major Soil Bacterial Phyla Communities and Physicochemical Factors.In the figure, “*” indicates P \u0026lt; 0.05 and ≥ 0.01, “**” indicates P \u0026lt; 0.01 and ≥ 0.005, and “***” indicates P \u0026lt; 0.005. The size of the square represents the magnitude of |r|. The same applies below.\u003c/p\u003e","description":"","filename":"image6.png","url":"https://assets-eu.researchsquare.com/files/rs-8998912/v1/f178c37881484969a3030c8f.png"},{"id":106230663,"identity":"29d6cd2d-a317-4112-85fb-cc4611159bfe","added_by":"auto","created_at":"2026-04-06 12:32:49","extension":"png","order_by":7,"title":"Figure 7","display":"","copyAsset":false,"role":"figure","size":117970,"visible":true,"origin":"","legend":"\u003cp\u003eCorrelation Analysis Between Major Fungal Phyla Communities and Physicochemical Factors in Soil\u003c/p\u003e","description":"","filename":"image7.png","url":"https://assets-eu.researchsquare.com/files/rs-8998912/v1/7c2e5bb58b49e490d00aac3c.png"},{"id":106403066,"identity":"2c93d32f-8431-43ee-9654-740c69799cbe","added_by":"auto","created_at":"2026-04-08 09:13:30","extension":"png","order_by":8,"title":"Figure 8","display":"","copyAsset":false,"role":"figure","size":101646,"visible":true,"origin":"","legend":"\u003cp\u003eEffects of Mealworm Castings Application on Soil PLS-SEM\u003c/p\u003e","description":"","filename":"image8.png","url":"https://assets-eu.researchsquare.com/files/rs-8998912/v1/aa4a99d2a07eabe1e4eab6b1.png"},{"id":106403216,"identity":"e10e1917-819b-4110-a4aa-a3df679ea71f","added_by":"auto","created_at":"2026-04-08 09:13:52","extension":"png","order_by":9,"title":"Figure 9","display":"","copyAsset":false,"role":"figure","size":97220,"visible":true,"origin":"","legend":"\u003cp\u003ePLS-SEM Analysis of How Soil Texture Modulates the Effects of Yellow Mealworm Castings on Soil.Data in the model were processed and calculated to determine the rate of change for each indicator between treatments with mealworm castings and their respective blank controls.\u003c/p\u003e","description":"","filename":"image9.png","url":"https://assets-eu.researchsquare.com/files/rs-8998912/v1/7fbca014f8c312e6a6617941.png"},{"id":107863765,"identity":"ebbfa0c2-9563-4239-a0af-67a39198aaea","added_by":"auto","created_at":"2026-04-27 06:11:53","extension":"pdf","order_by":0,"title":"","display":"","copyAsset":false,"role":"manuscript-pdf","size":1470586,"visible":true,"origin":"","legend":"","description":"","filename":"manuscript.pdf","url":"https://assets-eu.researchsquare.com/files/rs-8998912/v1/418721e3-1dac-4481-bb94-6e107ee29908.pdf"}],"financialInterests":"No competing interests reported.","formattedTitle":"Soil Mechanical Composition and Moisture Enhanced the Improvement of Mealworm Casting Amendment on Different Land Use Soils","fulltext":[{"header":"1 Introduction","content":"\u003cp\u003e \u003cem\u003eTenebrio molitor\u003c/em\u003e is a high-quality protein source and can be used as a premium feed additive for livestock or as a protein supplement in certain foods. Owing to its considerable economic value, it is widely cultivated on a large scale. As a holometabolous insect, the yellow mealworm is primarily commercialized at the larval stage before pupation. Large-scale larval rearing generates substantial quantities of mealworm castings (frass). Previous reports indicate that the ratio of \u003cem\u003eTenebrio molitor\u003c/em\u003e castings to dry \u003cem\u003eTenebrio molitor\u003c/em\u003e biomass is approximately 8.8:1. In 2021, the European Food Safety Authority (EFSA) formally authorized \u003cem\u003eTenebrio molitor\u003c/em\u003e as a novel food for human consumption. In China, \u003cem\u003eTenebrio molitor\u003c/em\u003e production reached 61,600 tons in 2023, while demand was 53,400 tons, and both figures continue to increase annually. Therefore, the efficient and scientifically sound utilization of mealworm castings in agricultural production to maximize their value remains an urgent issue that needs to be addressed.\u003c/p\u003e \u003cp\u003eMealworm castings are byproducts of yellow mealworm farming. They are typically described as dry, odorless, and sand-like, with a fine granular structure, and are easy to store and transport (Gao Yan, 2012; Shen, X. K. \u003cem\u003eet al\u003c/em\u003e., 2009). Studies have shown that mealworm castings are rich in organic matter and essential plant nutrients, including nitrogen, phosphorus, and potassium, and also contain insect-derived metabolites such as chitin and antimicrobial peptides. Compared with traditional livestock manure, mealworm castings have a more balanced nitrogen-to-phosphorus ratio and lower heavy metal contents, making them more environmentally friendly. In addition, they can improve soil structure, increase water-holding capacity, and enhance soil fertility (He et al., \u003cspan citationid=\"CR1\" class=\"CitationRef\"\u003e2021\u003c/span\u003e; Zunzunegui et al., \u003cspan citationid=\"CR17\" class=\"CitationRef\"\u003e2024\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eCurrently, few studies have examined the differential effects of mealworm castings across soils from different land-use types. Variation in the effects of organic fertilizers among land-use types represents a complex ecological issue shaped by multiple factors, including soil physicochemical properties, microbial communities, vegetation, and management practices (Li \u003cem\u003eet al\u003c/em\u003e., 2021). Numerous studies have shown that organic fertilizers can exert markedly different effects on soil fertility, nutrient cycling, and ecosystem functioning across croplands, grasslands, and forest lands (Fang \u003cem\u003eet al\u003c/em\u003e., 2013; Tang et al., \u003cspan citationid=\"CR6\" class=\"CitationRef\"\u003e2020\u003c/span\u003e; Zajicova \u003cem\u003eet al\u003c/em\u003e., 2019). Soil texture\u0026mdash;specifically the proportions of sand, silt, and clay\u0026mdash;regulates the effectiveness of organic fertilizers and the extent of fertility improvement by influencing soil physical structure, water-holding capacity, nutrient adsorption and release, and microbial activity (Adeniji, A. \u003cem\u003eet al\u003c/em\u003e., 2021). In addition, the bioavailability of soil organic carbon (SOC) is closely linked to microbial activity and is mediated by particle-size fractionation and soil morphology (Xiao, S. et al., \u003cspan citationid=\"CR4\" class=\"CitationRef\"\u003e2024\u003c/span\u003e). Therefore, optimizing mealworm casting application strategies according to soil texture is essential for enhancing agricultural productivity, improving soil health, and promoting sustainable agriculture.\u003c/p\u003e \u003cp\u003eCroplands, as the primary land-use type receiving organic fertilizer inputs, are also expected to be a major target for large-scale application of mealworm castings. Moderate soil moisture (typically close to field capacity) promotes the mineralization of organic matter, thereby facilitating the release of nutrients such as nitrogen, phosphorus, and potassium from organic fertilizers for crop uptake (Du, T \u003cem\u003eet al\u003c/em\u003e., 2022). At the same time, soil moisture is a key environmental factor shaping microbial communities; changes in moisture can alter microbial community composition and diversity (Xing, Y et al., \u003cspan citationid=\"CR8\" class=\"CitationRef\"\u003e2024\u003c/span\u003e). Moreover, organic fertilizer inputs and moisture conditions can jointly influence the functional diversity of key soil microorganisms.\u003c/p\u003e \u003cp\u003eOrganic fertilizers can exert broad positive effects on soil environments and crop yields by improving soil physicochemical properties and reshaping microbial communities (Ma, G. et al., \u003cspan citationid=\"CR9\" class=\"CitationRef\"\u003e2023\u003c/span\u003e; Xing, Y et al., \u003cspan citationid=\"CR8\" class=\"CitationRef\"\u003e2024\u003c/span\u003e). However, their effectiveness varies across land-use practices and is largely determined by initial soil fertility, soil texture, management practices (e.g., water management), and ecosystem characteristics (Bajgai, Y. \u003cem\u003eet al\u003c/em\u003e., 2023; Li, Z. \u003cem\u003eet al\u003c/em\u003e., 2024). In this study, we investigated the effects of adding mealworm castings at 1% (w/w) to cropland soils representing three land-use types. We further analyzed how soil moisture conditions modulate the impacts of mealworm castings on soil physicochemical properties and microbial communities, thereby providing a theoretical basis for the scientific utilization of mealworm castings.\u003c/p\u003e"},{"header":"2 Materials and methods","content":"\u003cdiv id=\"Sec3\" class=\"Section2\"\u003e \u003ch2\u003e2.1 Test soil and mealworm castings\u003c/h2\u003e \u003cp\u003eTo fugure out the effect of mealworm castings on soils, three soils of cropland, forest land and grassland with different soil texture were selected for soil incubation experiment. The cropland soil was sampled from Binzhou City, Shandong Province (37.3025N,117.366E), with a land use type of irrigated farmland; the forest land soil was collected from Guangzhou City, Guangdong Province (23.930N,113.2120E), classified as forest land; the grassland soil was collected from Guangzhou City, Guangdong Province (23.936N,113.2043E), classified as other grassland. The collected soil samples were coarsely ground, sieved through a 2 mm mesh, and naturally air-dried. Field water-holding capacity, particle size distribution, and moisture content were then measured. Land use types were classified according to GB/T 21010\u0026thinsp;\u0026minus;\u0026thinsp;2007. Basic physicochemical properties of soil and mealworm castings are detailed in Tables\u0026nbsp;\u003cspan refid=\"Tab1\" class=\"InternalRef\"\u003e1\u003c/span\u003e. Based on the International Soil Classification Triangle, all three soil textures were classified as sandy loam. The mealworm castings used in this experiment were obtained from \u003cem\u003eTenebrio molitor\u003c/em\u003e larvae reared with wheat bran in the laboratory. After collecting, the castings were sieved through a 60-mesh screen.\u003c/p\u003e \u003cp\u003e \u003cdiv class=\"gridtable\"\u003e\u003ctable float=\"Yes\" id=\"Tab1\" border=\"1\"\u003e \u003ccaption language=\"En\"\u003e \u003cdiv class=\"CaptionNumber\"\u003eTable 1\u003c/div\u003e \u003cdiv class=\"CaptionContent\"\u003e \u003cp\u003e.Basic Physical and Chemical Properties of Test Soils\u003c/p\u003e \u003c/div\u003e \u003c/caption\u003e \u003ccolgroup cols=\"5\"\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c1\" colnum=\"1\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c2\" colnum=\"2\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c3\" colnum=\"3\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c4\" colnum=\"4\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c5\" colnum=\"5\"\u003e\u003c/div\u003e \u003cthead\u003e \u003ctr\u003e \u003cth align=\"left\" colname=\"c1\"\u003e \u003cp\u003eSoil sample\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c2\"\u003e \u003cp\u003eM\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c3\"\u003e \u003cp\u003eDL\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c4\"\u003e \u003cp\u003eLN\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c5\"\u003e \u003cp\u003eFT\u003c/p\u003e \u003c/th\u003e \u003c/tr\u003e \u003c/thead\u003e \u003ctbody\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003epH\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e6.40\u0026thinsp;\u0026plusmn;\u0026thinsp;0.03\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e7.69\u0026thinsp;\u0026plusmn;\u0026thinsp;0.00\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e7.63\u0026thinsp;\u0026plusmn;\u0026thinsp;0.00\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e4.86\u0026thinsp;\u0026plusmn;\u0026thinsp;0.00\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eTC/(g\u0026middot;kg\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e395.0\u0026thinsp;\u0026plusmn;\u0026thinsp;2.4\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e21.1\u0026thinsp;\u0026plusmn;\u0026thinsp;0.1\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e6.3\u0026thinsp;\u0026plusmn;\u0026thinsp;0.1\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e23.8\u0026thinsp;\u0026plusmn;\u0026thinsp;0.1\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eTN/(g\u0026middot;kg\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e58.20\u0026thinsp;\u0026plusmn;\u0026thinsp;0.43\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e1.67\u0026thinsp;\u0026plusmn;\u0026thinsp;0.06\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e0.67\u0026thinsp;\u0026plusmn;\u0026thinsp;0.01\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e1.37\u0026thinsp;\u0026plusmn;\u0026thinsp;0.30\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eGravel content %\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e-\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e65.8\u0026thinsp;\u0026plusmn;\u0026thinsp;0.0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e72.1\u0026thinsp;\u0026plusmn;\u0026thinsp;0.0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e76.7\u0026thinsp;\u0026plusmn;\u0026thinsp;0.0\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eParticle content %\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e-\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e25.2\u0026thinsp;\u0026plusmn;\u0026thinsp;0.0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e26.9\u0026thinsp;\u0026plusmn;\u0026thinsp;0.0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e14.1\u0026thinsp;\u0026plusmn;\u0026thinsp;0.0\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eSilt content %\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e-\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e9.03\u0026thinsp;\u0026plusmn;\u0026thinsp;0.00\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e1.01\u0026thinsp;\u0026plusmn;\u0026thinsp;0.00\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e9.21\u0026thinsp;\u0026plusmn;\u0026thinsp;0.00\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eField capacity %\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e-\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e36.8\u0026thinsp;\u0026plusmn;\u0026thinsp;0.0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e48.3\u0026thinsp;\u0026plusmn;\u0026thinsp;0.0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e37.8\u0026thinsp;\u0026plusmn;\u0026thinsp;0.0\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eAir dried soil Moisture content %\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e-\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e1.87\u0026thinsp;\u0026plusmn;\u0026thinsp;0.00\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e1.96\u0026thinsp;\u0026plusmn;\u0026thinsp;0.00\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e1.12\u0026thinsp;\u0026plusmn;\u0026thinsp;0.00\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eTSC/(mg\u0026middot;kg\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e17820.7\u0026thinsp;\u0026plusmn;\u0026thinsp;1979.6\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e138.1\u0026thinsp;\u0026plusmn;\u0026thinsp;16.3\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e28.1\u0026thinsp;\u0026plusmn;\u0026thinsp;2.6\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e135.0\u0026thinsp;\u0026plusmn;\u0026thinsp;20.6\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eTSOC/(mg\u0026middot;kg\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e16496.6\u0026thinsp;\u0026plusmn;\u0026thinsp;1431.6\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e111.6\u0026thinsp;\u0026plusmn;\u0026thinsp;5.3\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e26.1\u0026thinsp;\u0026plusmn;\u0026thinsp;0.7\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e133.0\u0026thinsp;\u0026plusmn;\u0026thinsp;9.5\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eAvbN/(mg\u0026middot;kg\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e-\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e78.2\u0026thinsp;\u0026plusmn;\u0026thinsp;1.7\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e18.4\u0026thinsp;\u0026plusmn;\u0026thinsp;0.6\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e102.3\u0026thinsp;\u0026plusmn;\u0026thinsp;0.8\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eNH\u003csub\u003e4\u003c/sub\u003e\u003csup\u003e+\u003c/sup\u003e-N/(mg\u0026middot;kg\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e-\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e9.85\u0026thinsp;\u0026plusmn;\u0026thinsp;1.14\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e3.45\u0026thinsp;\u0026plusmn;\u0026thinsp;0.64\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e26.77\u0026thinsp;\u0026plusmn;\u0026thinsp;4.24\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eNO\u003csub\u003e3\u003c/sub\u003e\u003csup\u003e\u0026minus;\u003c/sup\u003e-N/(mg\u0026middot;kg\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e-\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e50.97\u0026thinsp;\u0026plusmn;\u0026thinsp;10.68\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e4.71\u0026thinsp;\u0026plusmn;\u0026thinsp;1.57\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e19.23\u0026thinsp;\u0026plusmn;\u0026thinsp;4.84\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eAvbP/(mg\u0026middot;kg\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e-\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e70.49\u0026thinsp;\u0026plusmn;\u0026thinsp;9.20\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e5.96\u0026thinsp;\u0026plusmn;\u0026thinsp;3.43\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e4.71\u0026thinsp;\u0026plusmn;\u0026thinsp;0.91\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eAvbK/(mg\u0026middot;kg\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e-\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e113.2\u0026thinsp;\u0026plusmn;\u0026thinsp;0.6\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e31.9\u0026thinsp;\u0026plusmn;\u0026thinsp;0.0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e29.4\u0026thinsp;\u0026plusmn;\u0026thinsp;3.6\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colspan=\"5\" nameend=\"c5\" namest=\"c1\"\u003e \u003cp\u003epH, soil acidity/alkalinity (soil-water ratio 1:2.5); TN, total nitrogen; TC, total carbon; TSC, soluble carbon; TSOC, soluble organic carbon; AvbN, alkali-hydrolyzable nitrogen; NH\u003csub\u003e4\u003c/sub\u003e\u003csup\u003e+\u003c/sup\u003e-N, ammonium; NO\u003csub\u003e3\u003c/sub\u003e\u003csup\u003e\u0026minus;\u003c/sup\u003e-N, nitrogen; AvbP, available phosphorus; AvbK, available potassium. Table values are presented as mean\u0026thinsp;\u0026plusmn;\u0026thinsp;standard deviation. M \u003cem\u003eTenebrio molitor\u003c/em\u003e castings; DL denotes croplands; LN denotes grassland; FT denotes forest land.\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003c/tbody\u003e \u003c/colgroup\u003e \u003c/table\u003e\u003c/div\u003e \u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec4\" class=\"Section2\"\u003e \u003ch2\u003e2.2 Experimental Design\u003c/h2\u003e \u003cp\u003eEight treatments were established in this experiment (see Table\u0026nbsp;\u003cspan refid=\"Tab2\" class=\"InternalRef\"\u003e2\u003c/span\u003e for details), with 3 replicates per treatment. The specific experimental procedure is as follows: Each beaker contained 100 g of dried soil (converted to air-dried soil based on moisture content). Samples were divided into separate 200 ml beakers according to treatment. Neatly arrange the beakers containing the divided samples in a plastic box. Place water at the bottom of the box for humidity maintenance, cover the box, and place it in a constant-temperature incubator. Incubate at 26\u0026deg;C. Every three days, add an appropriate amount of ultrapure water to each treatment sample to restore it to its original weight.\u003c/p\u003e \u003cp\u003eDestructive sampling was performed on days 14 and 28, followed by post-treatment preservation. A portion of samples was air-dried for storage; another portion was stored at 4\u0026deg;C for basic physicochemical parameter analysis; and a third portion was preserved at -80\u0026deg;C for microbial parameter analysis.\u003c/p\u003e \u003cp\u003e \u003cdiv class=\"gridtable\"\u003e\u003ctable float=\"Yes\" id=\"Tab2\" border=\"1\"\u003e \u003ccaption language=\"En\"\u003e \u003cdiv class=\"CaptionNumber\"\u003eTable 2\u003c/div\u003e \u003cdiv class=\"CaptionContent\"\u003e \u003cp\u003e Basic Information for Each Treatment\u003c/p\u003e \u003c/div\u003e \u003c/caption\u003e \u003ccolgroup cols=\"4\"\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c1\" colnum=\"1\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c2\" colnum=\"2\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c3\" colnum=\"3\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c4\" colnum=\"4\"\u003e\u003c/div\u003e \u003cthead\u003e \u003ctr\u003e \u003cth align=\"left\" colname=\"c1\"\u003e \u003cp\u003eProcessing Number\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c2\"\u003e \u003cp\u003eLand Use Type\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c3\"\u003e \u003cp\u003eSoil Type Code\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c4\"\u003e \u003cp\u003eProcessing Method\u003c/p\u003e \u003c/th\u003e \u003c/tr\u003e \u003c/thead\u003e \u003ctbody\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eD6M\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eCropland\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eDL\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eAdd 1% mealworm castings (based on 1% of the dry weight of the soil sample) at 60% of the field capacity.\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eD6\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eCropland\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eDL\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e60% of the field capacity\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eD8M\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eCropland\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eDL\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eAdd 1% mealworm castings (based on 1% of the dry weight of the soil sample) at 80% of the field capacity.\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eD8\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eCropland\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eDL\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e80% of the field capacity\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eL6M\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eGrassland\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eLN\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eAdd 1% mealworm castings (based on 1% of the dry weight of the soil sample) at 60% of the field capacity.\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eL6\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eGrassland\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eLN\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e60% of the field capacity\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eF6M\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eForest land\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eFT\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eAdd 1% mealworm castings (based on 1% of the dry weight of the soil sample) at 60% of the field capacity.\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eF6\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eForest land\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eFT\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e60% of the field capacity\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003c/tbody\u003e \u003c/colgroup\u003e \u003c/table\u003e\u003c/div\u003e \u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec5\" class=\"Section2\"\u003e \u003ch2\u003e2.3 Soil Physicochemical Properties Testing\u003c/h2\u003e \u003cp\u003eThe determination of soil physicochemical parameters followed Bao Shidan's Soil Agrochemical Analysis (China Agricultural Press, 2000). Soil pH (soil : water\u0026thinsp;=\u0026thinsp;1 : 2.5) was measured using the potentiometric method; soil electrical conductivity was determined under extraction conditions consistent with soil pH, followed by measurement with a conductivity meter; soil available phosphorus content was determined using the ammonium fluoride extraction method; soil available potassium was determined by flame photometry; soil total nitrogen and total carbon were measured using a fully automated elemental analyzer (Manufacturer: Elementar, Model: Vario Micro cube); soil soluble organic carbon and soluble total nitrogen were determined by water extraction using a total organic carbon analyzer (Manufacturer: Elementar, Model: Vario TOC); Soil ammonium nitrogen was determined using the indophenol blue colorimetric method; Soil nitrate nitrogen was determined using the potassium chloride extraction method; Alkali-hydrolyzable nitrogen was determined using boric acid absorption titration.\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec6\" class=\"Section2\"\u003e \u003ch2\u003e2.4 Soil Microbial Sequencing\u003c/h2\u003e \u003cp\u003eMicrobial community analysis for this experiment was conducted by Beijing Novogene Co., Ltd., which performed extraction and sequencing using the Illumina NovaSeq PE250 platform. Fungi were sequenced using ITS amplicon sequencing, targeting the ITS1-1F region with primer sequences: ITS5-1737 F (5\u0026rsquo;-CTTGGTCATTTAGAGGAAGTAA-3\u0026rsquo;) and ITS2-2043 R (5\u0026rsquo;-GCTGCGTTCTTCATCGATGC-3\u0026rsquo;). Bacteria were sequenced using the 16S rDNA amplicon, with the amplification region set to 16S V3-V4. Primer sequences were: 515 F (5\u0026rsquo;-CCTAYGGGRBGCASCAG-3\u0026rsquo;) and 806 R (5\u0026rsquo;-GGACTACNNGGGTATCTAAT-3\u0026rsquo;).\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec7\" class=\"Section2\"\u003e \u003ch2\u003e1.5 Data Analysis and Statistical Methods\u003c/h2\u003e \u003cp\u003eData analysis and processing were performed using Microsoft Office Excel 2013 (Microsoft Corporation, USA), SPSS 18.0 (IBM Corporation), and RStudio (RStudio Company). Alpha diversity and Beta diversity analyses were conducted using the R software version 4.2.3. The raw sequencing data (Raw Data) contained a certain proportion of interfering data (Dirty Data). To ensure that the results of the information analysis were more accurate and reliable, the raw data were first merged and filtered to obtain effective data (Clean Data). Then, the effective data were denoised using DADA2 or deblur (Li M et al., \u003cspan citationid=\"CR24\" class=\"CitationRef\"\u003e2020\u003c/span\u003e), thereby yielding the final ASVs. For the obtained ASVs (Callahan BJ \u003cem\u003eet al\u003c/em\u003e., 2017), on the one hand, the representative sequence of each ASV was subjected to taxonomic annotation to obtain the corresponding species information and species-based abundance distribution. The classify-sklearn algorithm in QIIME2 (Bokulich NA et al., \u003cspan citationid=\"CR25\" class=\"CitationRef\"\u003e2018\u003c/span\u003e; Bolyen E \u003cem\u003eet al\u003c/em\u003e., 2019) was used to annotate each ASV using a pre-trained Naive Bayes classifier. The annotation database was Silva 138.1.\u003c/p\u003e \u003c/div\u003e"},{"header":"3 Results Analysis","content":"\u003cdiv id=\"Sec9\" class=\"Section2\"\u003e \u003ch2\u003e3.1 Effects of \u003cem\u003eTenebrio molitor\u003c/em\u003e Castings on Soil Physicochemical Properties\u003c/h2\u003e \u003cp\u003eAs shown in Table\u0026nbsp;\u003cspan refid=\"Tab3\" class=\"InternalRef\"\u003e3\u003c/span\u003e, compared with the blank control, the addition of mealworm castings significantly increased the content of soil total nitrogen (TN), soil soluble carbon (TSC), soil soluble organic carbon (TSOC), alkali-hydrolyzable nitrogen (AvbN), ammonium nitrogen (NH\u003csub\u003e4\u003c/sub\u003e\u003csup\u003e+\u003c/sup\u003e-N), and soil nitrate nitrogen (NO\u003csub\u003e3\u003c/sub\u003e\u003csup\u003e\u0026minus;\u003c/sup\u003e-N) in both forest (FT) and grassland (LN) soils. The increases ranged from 15.0% to 31.6%, 37.2% to 171.1%, 41.5% to 178.0%, 73.9% to 203.8%, 184.0% to 620.7%, and 31.5% to 581.9%, respectively. The addition of mealworm castings significantly increased the pH value of forest land by 0.3\u0026ndash;0.5 units. For grassland, the pH value decreased significantly by 0.2 units on day 28.\u003c/p\u003e \u003cp\u003eCompared to the blank control, insect dung addition also significantly increased available potassium (AvbK) and available phosphorus (AvbP) in forest land (FT), grassland (LN), and cropland (DL) (P\u0026thinsp;\u0026lt;\u0026thinsp;0.05), with increases ranging from 116% to 672% for AvbK and from 23.7% to 726.7% for AvbP.\u003c/p\u003e \u003cp\u003eAmong different cropland-capacity moisture treatments, adding mealworm castings had no significant effect on soil soluble carbon content in the 60% cropland-capacity treatment (DL6). However, it significantly increased soluble carbon content in the 80% cropland-capacity treatment (DL8) by 56.3% on day 28. Compared to the blank control, the DL6 treatment with added mealworm castings showed a significant increase in soil soluble organic carbon at day 14, while the DL8 treatment exhibited a significant increase only at day 28, with increases of 49.2% and 56.3%, respectively. Compared to the blank control, adding mealworm castings significantly increased total soil nitrogen in the DL6 treatment, with increases ranging from 11.3% to 11.7%. In the DL8 treatment, adding mealworm castings significantly reduced soil nitrate nitrogen content compared to the blank, with a decrease ranging from 73.2% to 87.4%. In DL treatments, ammonium nitrogen content significantly increased by 128%\u0026ndash;176% at day 28. In the DL8 treatment, soil ammonium nitrogen content increased significantly on day 14, with an increase of 152.3%. DL6 treatment with mealworm castings significantly lowered pH by 0.08\u0026ndash;0.16 units, while DL8 treatment with mealworm castings significantly raised pH by 0.08\u0026ndash;0.14 units.\u003c/p\u003e \u003cp\u003eAs shown in Fig.\u0026nbsp;\u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e1\u003c/span\u003e, PC1 explained 50.34% of the variance, PC2 explained 24.98%, and the cumulative explanation reached 75.32%. The treatments incorporating mealworm castings showed distinct differentiation from the blank control and original soil samples. Compared to the blank control, the addition of mealworm castings resulted in a clear differentiation along the PC1 axis overall. This indicates that adding mealworm castings significantly increased soil alkaline-hydrolyzable nitrogen content, soil soluble nitrogen content, soil soluble carbon content, soil total carbon content, soil total nitrogen content, ammonium nitrogen content, nitrate nitrogen content, available phosphorus content, and available potassium content. The treatment with added mealworm castings showed distinct differentiation on the PC2 axis. This indicates that over time, the addition of mealworm castings does not affect soil total carbon content, has a minor impact on total nitrogen content, but significantly influences pH and ammonium nitrogen content. Among treatments with different moisture contents, the treatment at 60% field capacity showed more pronounced differentiation on the PC1 axis compared to the treatment at 80% field capacity.\u003c/p\u003e \u003cp\u003e \u003cdiv class=\"gridtable\"\u003e\u003ctable float=\"Yes\" id=\"Tab3\" border=\"1\"\u003e \u003ccaption language=\"En\"\u003e \u003cdiv class=\"CaptionNumber\"\u003eTable 3\u003c/div\u003e \u003cdiv class=\"CaptionContent\"\u003e \u003cp\u003e Effects of Different Treatments on Soil Physicochemical Properties\u003c/p\u003e \u003c/div\u003e \u003c/caption\u003e \u003ccolgroup cols=\"11\"\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c1\" colnum=\"1\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c2\" colnum=\"2\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c3\" colnum=\"3\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c4\" colnum=\"4\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c5\" colnum=\"5\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c6\" colnum=\"6\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c7\" colnum=\"7\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c8\" colnum=\"8\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c9\" colnum=\"9\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c10\" colnum=\"10\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c11\" colnum=\"11\"\u003e\u003c/div\u003e \u003cthead\u003e \u003ctr\u003e \u003cth align=\"left\" colname=\"c1\"\u003e \u003cp\u003esample\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c2\"\u003e \u003cp\u003epH\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c3\"\u003e \u003cp\u003eTN\u003c/p\u003e \u003cp\u003e%\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c4\"\u003e \u003cp\u003eTC\u003c/p\u003e \u003cp\u003e%\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c5\"\u003e \u003cp\u003eTSC\u003c/p\u003e \u003cp\u003e/(mg\u0026middot;kg\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e)\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c6\"\u003e \u003cp\u003eTSOC\u003c/p\u003e \u003cp\u003e/(mg\u0026middot;kg\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e)\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c7\"\u003e \u003cp\u003eAvbN\u003c/p\u003e \u003cp\u003e/(mg\u0026middot;kg\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e)\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c8\"\u003e \u003cp\u003eNH\u003csub\u003e4\u003c/sub\u003e\u003csup\u003e+\u003c/sup\u003e-N\u003c/p\u003e \u003cp\u003e/(mg\u0026middot;kg\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e)\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c9\"\u003e \u003cp\u003eNO\u003csub\u003e3\u003c/sub\u003e\u003csup\u003e\u0026minus;\u003c/sup\u003e-N\u003c/p\u003e \u003cp\u003e/(mg\u0026middot;kg\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e)\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c10\"\u003e \u003cp\u003eAvbP\u003c/p\u003e \u003cp\u003e/(mg\u0026middot;kg\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e)\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c11\"\u003e \u003cp\u003eAvbK\u003c/p\u003e \u003cp\u003e/(mg\u0026middot;kg\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e)\u003c/p\u003e \u003c/th\u003e \u003c/tr\u003e \u003c/thead\u003e \u003ctbody\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cb\u003ed14D6M\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e7.35\u0026thinsp;\u0026plusmn;\u0026thinsp;0.02c\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" 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\u003cp\u003e8.80\u0026thinsp;\u0026plusmn;\u0026thinsp;0.70a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e34.5\u0026thinsp;\u0026plusmn;\u0026thinsp;0.8b\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e33.1\u0026thinsp;\u0026plusmn;\u0026thinsp;1.2b\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e53.8\u0026thinsp;\u0026plusmn;\u0026thinsp;1.7b\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e13.55\u0026thinsp;\u0026plusmn;\u0026thinsp;1.41b\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003e18.63\u0026thinsp;\u0026plusmn;\u0026thinsp;1.33a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003e12.7\u0026thinsp;\u0026plusmn;\u0026thinsp;2.3a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c11\"\u003e \u003cp\u003e198.3\u0026thinsp;\u0026plusmn;\u0026thinsp;4a\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cb\u003ed28L6\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e7.24\u0026thinsp;\u0026plusmn;\u0026thinsp;0.03a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e0.63\u0026thinsp;\u0026plusmn;\u0026thinsp;0.06b\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e6.83\u0026thinsp;\u0026plusmn;\u0026thinsp;0.31b\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e14.5\u0026thinsp;\u0026plusmn;\u0026thinsp;2.1c\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e13.4\u0026thinsp;\u0026plusmn;\u0026thinsp;0.7c\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e24.1\u0026thinsp;\u0026plusmn;\u0026thinsp;2.2c\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e4.77\u0026thinsp;\u0026plusmn;\u0026thinsp;0.75c\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003e2.73\u0026thinsp;\u0026plusmn;\u0026thinsp;0.43c\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003e3.7\u0026thinsp;\u0026plusmn;\u0026thinsp;2.2b\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c11\"\u003e \u003cp\u003e30.4\u0026thinsp;\u0026plusmn;\u0026thinsp;1.1b\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cb\u003ed28F6M\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e4.98\u0026thinsp;\u0026plusmn;\u0026thinsp;0.02b\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e1.57\u0026thinsp;\u0026plusmn;\u0026thinsp;0.15a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e21.17\u0026thinsp;\u0026plusmn;\u0026thinsp;0.61ab\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e118.8\u0026thinsp;\u0026plusmn;\u0026thinsp;29.0ab\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e115.1\u0026thinsp;\u0026plusmn;\u0026thinsp;26.6ab\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e157.0\u0026thinsp;\u0026plusmn;\u0026thinsp;8.0b\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e33.36\u0026thinsp;\u0026plusmn;\u0026thinsp;4.87b\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003e32.27\u0026thinsp;\u0026plusmn;\u0026thinsp;1.07a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003e28.6\u0026thinsp;\u0026plusmn;\u0026thinsp;3.1a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c11\"\u003e \u003cp\u003e195.7\u0026thinsp;\u0026plusmn;\u0026thinsp;2.9a\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cb\u003ed28F6\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e4.65\u0026thinsp;\u0026plusmn;\u0026thinsp;0.06c\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e1.50\u0026thinsp;\u0026plusmn;\u0026thinsp;0.10ab\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e22.70\u0026thinsp;\u0026plusmn;\u0026thinsp;1.04a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e63.1\u0026thinsp;\u0026plusmn;\u0026thinsp;10.4c\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e49.7\u0026thinsp;\u0026plusmn;\u0026thinsp;2.2c\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e90.3\u0026thinsp;\u0026plusmn;\u0026thinsp;0.8c\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e10.20\u0026thinsp;\u0026plusmn;\u0026thinsp;1.92c\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003e24.54\u0026thinsp;\u0026plusmn;\u0026thinsp;4.15ab\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003e4.9\u0026thinsp;\u0026plusmn;\u0026thinsp;1.1c\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c11\"\u003e \u003cp\u003e25.3\u0026thinsp;\u0026plusmn;\u0026thinsp;1.1b\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colspan=\"11\" nameend=\"c11\" namest=\"c1\"\u003e \u003cp\u003eValues in the table represent mean\u0026thinsp;\u0026plusmn;\u0026thinsp;standard error (n\u0026thinsp;=\u0026thinsp;3). Different lowercase letters indicate significant differences among treatments within the same land use type and moisture content (P\u0026thinsp;\u0026lt;\u0026thinsp;0.05). Treatment names: d14 denotes day 14 of cultivation, d28 denotes day 28; M indicates addition of mealworm castings, 6 denotes 60% field capacity moisture content, 8 denotes 80% field capacity moisture content; D denotes croplands, L denotes grassland, F denotes forest land. Among replicates, the coefficient of variation (CV; standard deviation/mean) was calculated. When the CV exceeded 0.3, one outlier was removed from the three replicate measurements to minimize the CV as much as possible.\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003c/tbody\u003e \u003c/colgroup\u003e \u003c/table\u003e\u003c/div\u003e \u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec10\" class=\"Section2\"\u003e \u003ch2\u003e3.2 Analysis of the Effects of \u003cem\u003eTenebrio molitor\u003c/em\u003e Castings on Soil Microbial Community Diversity\u003c/h2\u003e \u003cdiv id=\"Sec11\" class=\"Section3\"\u003e \u003ch2\u003e3.2.1 Alpha Analysis of Soil Microbial Communities\u003c/h2\u003e \u003cp\u003eAnalysis of diversity indices for the different treatments is shown in Table\u0026nbsp;\u003cspan refid=\"Tab4\" class=\"InternalRef\"\u003e4\u003c/span\u003e. When insect frass was added to different land-use types, the bacterial Shannon index decreased significantly in the cultivated land (DL6) and grassland (LN6) treatments compared with the blank control, with reductions ranging from 4.67% to 7.83%. Whereas the Shannon index for fungi showed a significant increase in the forest land (FT6) treatment, with the rate of increase intensifying over time, rising from 1.75% on day 14 to 5.98% on day 28. The variation in the Shannon index for fungi across different treatments was similar to that observed for bacteria; the decline in the DL treatments was relatively small, ranging from 12.1% to 23.6%; the LN treatment showed a larger decrease, at 30.6% on day 14 and 25.2% on day 28; the FT treatment showed a slight increase after the addition of insect faeces, but this was not statistically significant. Regarding the bacterial Simpson\u0026rsquo;s index, following the addition of insect faeces, only the cultivated land treatment showed a significant decrease compared with the blank control; the grassland treatment showed no significant change, whilst the woodland treatment showed a significant increase on day 28. The Simpson index for fungi showed more pronounced changes. Following the addition of insect faeces, the arable land treatment showed no significant change; the grassland treatment exhibited a significant decrease of 13.0% on day 14, which improved slightly by day 28 compared to day 14, yet the treatment with added insect faeces remained significantly lower than the blank control, with a decrease of 7.42%; The trend in the forest plot was the opposite to that of the other treatments; the fungal Simpson\u0026rsquo;s index in the insect faeces-added treatment was significantly higher than that of the blank control, with an increase of 33.4%\u0026ndash;44.3%. The changes in the bacterial ACE and Chao-1 indices were consistent: following the addition of insect faeces, the arable land treatment showed a significant decrease compared to the blank control; the grassland treatment also showed a significant decrease compared to the blank control; however, there was no significant difference among the forest plot treatments. The trends in fungal ACE and Chao-1 indices were consistent: following the addition of insect frass, the arable land, forest land and grassland treatments all showed a significant decrease compared to the control, with a reduction ranging from 21.6% to 57.3%.\u003c/p\u003e \u003cp\u003eTranslated In cultivated land, the effects of different moisture content treatments on microbial diversity also varied considerably. In the treatment with a moisture content of 80% of field capacity (DL8), the treatment supplemented with insect frass showed, on day 14, that the bacterial Shannon, bacterial Simpson, bacterial ACE, bacterial Chao1, the fungal ACE and Chao-1 indices were significantly lower than those of the control treatment; however, by day 28, only the fungal ACE and Chao-1 indices were significantly lower than those of the control treatment; whereas there were no significant differences between treatments in the fungal Simpson and Shannon indices.\u003c/p\u003e \u003cp\u003e \u003cdiv class=\"gridtable\"\u003e\u003ctable float=\"Yes\" id=\"Tab4\" border=\"1\"\u003e \u003ccaption language=\"En\"\u003e \u003cdiv class=\"CaptionNumber\"\u003eTable 4\u003c/div\u003e \u003cdiv class=\"CaptionContent\"\u003e \u003cp\u003e Assessment of Soil Microbial Diversity Under Different Treatments\u003c/p\u003e \u003c/div\u003e \u003c/caption\u003e \u003ccolgroup cols=\"9\"\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c1\" colnum=\"1\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c2\" colnum=\"2\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c3\" colnum=\"3\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c4\" colnum=\"4\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c5\" colnum=\"5\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c6\" colnum=\"6\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c7\" colnum=\"7\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c8\" colnum=\"8\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c9\" colnum=\"9\"\u003e\u003c/div\u003e \u003cthead\u003e \u003ctr\u003e \u003cth align=\"left\" colname=\"c1\"\u003e \u003cp\u003eTreatments\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c2\"\u003e \u003cp\u003eBacterial Shannon Index\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c3\"\u003e \u003cp\u003eBacterial Simpson Index\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c4\"\u003e \u003cp\u003eBacterial\u003c/p\u003e \u003cp\u003eChao 1 Index\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c5\"\u003e \u003cp\u003eBacterial\u003c/p\u003e \u003cp\u003eACE Index\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c6\"\u003e \u003cp\u003eFungal Shannon Index\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c7\"\u003e \u003cp\u003eFungal Simpson Index\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c8\"\u003e \u003cp\u003eFungal\u003c/p\u003e \u003cp\u003eChao 1 Index\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c9\"\u003e \u003cp\u003eFungal\u003c/p\u003e \u003cp\u003eACE Index\u003c/p\u003e \u003c/th\u003e \u003c/tr\u003e \u003c/thead\u003e \u003ctbody\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cb\u003ed0DL\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e6.81\u0026thinsp;\u0026plusmn;\u0026thinsp;0.06a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e1.00\u0026thinsp;\u0026plusmn;\u0026thinsp;0.00a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e2256\u0026thinsp;\u0026plusmn;\u0026thinsp;35a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e2256\u0026thinsp;\u0026plusmn;\u0026thinsp;35a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e3.27\u0026thinsp;\u0026plusmn;\u0026thinsp;0.09a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e0.90\u0026thinsp;\u0026plusmn;\u0026thinsp;0.01a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e318\u0026thinsp;\u0026plusmn;\u0026thinsp;24ab\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003e318\u0026thinsp;\u0026plusmn;\u0026thinsp;24ab\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cb\u003ed0LN\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e7.20\u0026thinsp;\u0026plusmn;\u0026thinsp;0.08a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e1.00\u0026thinsp;\u0026plusmn;\u0026thinsp;0.00a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e2583\u0026thinsp;\u0026plusmn;\u0026thinsp;142a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e2583\u0026thinsp;\u0026plusmn;\u0026thinsp;142a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e4.38\u0026thinsp;\u0026plusmn;\u0026thinsp;0.08a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e0.96\u0026thinsp;\u0026plusmn;\u0026thinsp;0a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e515\u0026thinsp;\u0026plusmn;\u0026thinsp;22a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003e515\u0026thinsp;\u0026plusmn;\u0026thinsp;22a\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cb\u003ed0FT\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e6.63\u0026thinsp;\u0026plusmn;\u0026thinsp;0.11a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e1.00\u0026thinsp;\u0026plusmn;\u0026thinsp;0.00ab\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e1872\u0026thinsp;\u0026plusmn;\u0026thinsp;403a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e1872\u0026thinsp;\u0026plusmn;\u0026thinsp;403a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e2.36\u0026thinsp;\u0026plusmn;\u0026thinsp;0.22a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e0.70\u0026thinsp;\u0026plusmn;\u0026thinsp;0.05a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e286\u0026thinsp;\u0026plusmn;\u0026thinsp;27a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003e286\u0026thinsp;\u0026plusmn;\u0026thinsp;27a\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cb\u003ed14DL60M\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e6.10\u0026thinsp;\u0026plusmn;\u0026thinsp;0.1c\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e0.99\u0026thinsp;\u0026plusmn;\u0026thinsp;0.00cd\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e1472\u0026thinsp;\u0026plusmn;\u0026thinsp;105c\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e1472\u0026thinsp;\u0026plusmn;\u0026thinsp;105c\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e2.51\u0026thinsp;\u0026plusmn;\u0026thinsp;0.2b\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e0.82\u0026thinsp;\u0026plusmn;\u0026thinsp;0.03a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e191\u0026thinsp;\u0026plusmn;\u0026thinsp;19c\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e 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\u003cp\u003e1.00\u0026thinsp;\u0026plusmn;\u0026thinsp;0.00ab\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e1505\u0026thinsp;\u0026plusmn;\u0026thinsp;87bc\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e1505\u0026thinsp;\u0026plusmn;\u0026thinsp;87bc\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e2.65\u0026thinsp;\u0026plusmn;\u0026thinsp;0.43bc\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e0.81\u0026thinsp;\u0026plusmn;\u0026thinsp;0.07b\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e245\u0026thinsp;\u0026plusmn;\u0026thinsp;54bc\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003e245\u0026thinsp;\u0026plusmn;\u0026thinsp;54bc\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cb\u003ed28DL80\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e6.45\u0026thinsp;\u0026plusmn;\u0026thinsp;0.16b\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e1.00\u0026thinsp;\u0026plusmn;\u0026thinsp;0.00a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e1616\u0026thinsp;\u0026plusmn;\u0026thinsp;123b\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e1616\u0026thinsp;\u0026plusmn;\u0026thinsp;123b\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e3.02\u0026thinsp;\u0026plusmn;\u0026thinsp;0.09ab\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e0.83\u0026thinsp;\u0026plusmn;\u0026thinsp;0.02ab\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e322\u0026thinsp;\u0026plusmn;\u0026thinsp;50a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003e322\u0026thinsp;\u0026plusmn;\u0026thinsp;50a\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cb\u003ed28LN60M\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e6.31\u0026thinsp;\u0026plusmn;\u0026thinsp;0.13c\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e0.99\u0026thinsp;\u0026plusmn;\u0026thinsp;0.00bc\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e1765\u0026thinsp;\u0026plusmn;\u0026thinsp;128cd\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e1765\u0026thinsp;\u0026plusmn;\u0026thinsp;128cd\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e3.31\u0026thinsp;\u0026plusmn;\u0026thinsp;0.42c\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e0.90\u0026thinsp;\u0026plusmn;\u0026thinsp;0.05b\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e181\u0026thinsp;\u0026plusmn;\u0026thinsp;4d\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003e181\u0026thinsp;\u0026plusmn;\u0026thinsp;4d\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cb\u003ed28LN60\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e6.75\u0026thinsp;\u0026plusmn;\u0026thinsp;0.04b\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e1.00\u0026thinsp;\u0026plusmn;\u0026thinsp;0.00b\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e2176\u0026thinsp;\u0026plusmn;\u0026thinsp;128b\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e2176\u0026thinsp;\u0026plusmn;\u0026thinsp;128b\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e4.42\u0026thinsp;\u0026plusmn;\u0026thinsp;0.11a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e0.97\u0026thinsp;\u0026plusmn;\u0026thinsp;0.01a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e424\u0026thinsp;\u0026plusmn;\u0026thinsp;4b\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003e424\u0026thinsp;\u0026plusmn;\u0026thinsp;4b\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cb\u003ed28FT60M\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e6.60\u0026thinsp;\u0026plusmn;\u0026thinsp;0.17a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e1.00\u0026thinsp;\u0026plusmn;\u0026thinsp;0.00a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e1772\u0026thinsp;\u0026plusmn;\u0026thinsp;265ab\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e1772\u0026thinsp;\u0026plusmn;\u0026thinsp;265ab\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e1.72\u0026thinsp;\u0026plusmn;\u0026thinsp;0.16b\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e0.65\u0026thinsp;\u0026plusmn;\u0026thinsp;0.05a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e191\u0026thinsp;\u0026plusmn;\u0026thinsp;13b\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003e191\u0026thinsp;\u0026plusmn;\u0026thinsp;13b\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cb\u003ed28FT60\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e6.23\u0026thinsp;\u0026plusmn;\u0026thinsp;0.17b\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e0.99\u0026thinsp;\u0026plusmn;\u0026thinsp;0.00ab\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e1446\u0026thinsp;\u0026plusmn;\u0026thinsp;110ab\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e1446\u0026thinsp;\u0026plusmn;\u0026thinsp;110ab\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e1.49\u0026thinsp;\u0026plusmn;\u0026thinsp;0.39b\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e0.45\u0026thinsp;\u0026plusmn;\u0026thinsp;0.12b\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e273\u0026thinsp;\u0026plusmn;\u0026thinsp;26a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003e273\u0026thinsp;\u0026plusmn;\u0026thinsp;26a\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colspan=\"9\" nameend=\"c9\" namest=\"c1\"\u003e \u003cp\u003eValues in the table represent mean\u0026thinsp;\u0026plusmn;\u0026thinsp;standard error (n\u0026thinsp;=\u0026thinsp;3). Different lowercase letters indicate significant differences among treatments within the same land use type and moisture content (P\u0026thinsp;\u0026lt;\u0026thinsp;0.05). Treatment designations: d14 denotes day 14 of cultivation, d28 denotes day 28; M indicates addition of mealworm castings, 6 denotes 60% field capacity moisture content, 8 denotes 80% field capacity moisture content; D denotes croplands, L denotes grassland, F denotes forest land.\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003c/tbody\u003e \u003c/colgroup\u003e \u003c/table\u003e\u003c/div\u003e \u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec12\" class=\"Section3\"\u003e \u003ch2\u003e3.2.2 Beta Analysis of Soil Microbial Communities\u003c/h2\u003e \u003cp\u003eThe RDA analysis of bacterial communities is shown in Fig.\u0026nbsp;\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e2\u003c/span\u003e(a). RDA1 explained 46.34% of the variation, RDA2 explained 32.17%, and the cumulative explanation reached 78.51%. The physicochemical factors pH, available phosphorus (AvbP), and available potassium (AvbK) had the greatest effects on the bacterial community. Overall, the bacterial community composition in soils amended with insect frass was clearly differentiated from that in soils without insect frass addition. In the grassland (LN) and forest land (FT) treatments, the bacterial communities in the insect frass-amended treatments were clearly differentiated from those in the blank controls; the differences between treatments were more pronounced on day 14 and became smaller on day 28. In the cropland treatment (DL), the treatment with 60% field water-holding capacity (DL 6) showed clear differentiation from the blank control after insect frass addition, but the degree of differentiation was smaller than that in the grassland and forest land treatments. Compared with the blank control, the treatment with 80% field water-holding capacity (DL 8) showed relatively clear differentiation on day 14 after insect frass addition, whereas the differentiation was not obvious on day 28. Among the three different land-use types, the forest land treatment showed the most pronounced difference from the blank control after insect frass addition.\u003c/p\u003e \u003cp\u003eRDA analysis of fungal communities is shown in Fig.\u0026nbsp;\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e2\u003c/span\u003e(b). RDA1 explained 85.68% of the variation, RDA2 explained 9.41%, and the cumulative explanation reached 95.09%; the explanatory power of the RDA1 axis was much greater than that of the RDA2 axis. Among the physicochemical factors, pH had the greatest effect on the fungal community, followed by available phosphorus and available potassium. Compared with the blank control, all treatments amended with insect frass showed clear differentiation along the RDA2 axis, indicating obvious differences in fungal communities. In the grassland treatments, compared with the blank control, the differences between the insect frass-amended treatments were more pronounced on day 14 and became smaller on day 28. In the forest land treatments, compared with the blank control, the differences in fungal communities in the insect frass-amended treatments became increasingly greater over time. In the cropland treatments, for the treatment with 60% field water-holding capacity, the difference from the blank control did not decrease obviously over time after insect frass addition. Compared with the DL 6 treatment, the treatment with 80% field water-holding capacity showed relatively small differences in fungal communities after insect frass addition. In addition to pH, available phosphorus, and available potassium, total nitrogen, total carbon, and soil soluble carbon also had relatively large effects on fungal community composition.\u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003c/div\u003e \u003c/div\u003e \u003cdiv id=\"Sec13\" class=\"Section2\"\u003e \u003ch2\u003e3.3 Effects of \u003cem\u003eTenebrio molitor\u003c/em\u003e Castings on Soil Microbial Community Structure and Composition\u003c/h2\u003e \u003cp\u003eIn this study, we performed ASV (Amplicon Sequence Variant) clustering analysis on the valid sequences from all samples. Following species annotation, the top 10 bacterial phyla were selected for analysis, as shown in Fig.\u0026nbsp;\u003cspan refid=\"Fig3\" class=\"InternalRef\"\u003e3\u003c/span\u003e(a). The results showed that the bacterial communities exhibited clear differences at the phylum level under different soil types and incubation treatments, and that incubation time, moisture content, and insect frass addition had different effects on the relative abundances of the dominant bacterial phyla. Overall, the dominant bacterial phyla in samples from all treatments mainly included Pseudomonadota, Actinomycetota, Acidobacteriota, Bacteroidota, Bacillota, and Gemmatimonadota, but the variation trends were not consistent among different soils. In the cropland treatments, the insect frass-amended treatment at 60% field water-holding capacity (DL 6) showed high abundances of Pseudomonadota and Bacteroidota on days 14 and 28. Over time from day 14 to day 28, the abundance of Gemmatimonadota further increased under insect frass addition. In the high-moisture treatment (DL 8) amended with insect frass, the abundances of Pseudomonadota, Bacteroidota, and Gemmatimonadota were high on days 14 and 28. However, on day 28, the abundance of Pseudomonadota decreased, whereas that of Bacillota increased under insect frass addition. Over time from day 14 to day 28, Pseudomonadota showed a decreasing trend, whereas Gemmatimonadota showed an increasing trend under insect frass addition. In the grassland (LN) and forest land (FT) treatments, the 60% field water-holding capacity treatment amended with insect frass showed that Pseudomonadota and Acidobacteriota maintained high abundances on days 14 and 28. Over time from day 14 to day 28, the abundance of Acidobacteriota further increased under insect frass addition.\u003c/p\u003e \u003cp\u003eAnalysis of the top 10 fungal phyla at the phylum level is shown in Fig.\u0026nbsp;\u003cspan refid=\"Fig3\" class=\"InternalRef\"\u003e3\u003c/span\u003e(b). In the cropland treatments, the insect frass-amended treatment at 60% field water-holding capacity (DL 6) showed a relatively high abundance of Ascomycota on day 14. On day 28, Ascomycota still maintained a high abundance under insect frass addition, whereas Basidiomycota and Others were relatively higher in the control treatment. Over time from day 14 to day 28, the major fungal phyla showed little overall change under insect frass addition. In the high-moisture treatment (DL 8) amended with insect frass, Ascomycota showed a high abundance on day 14, whereas Others showed a relatively low abundance. On day 28, Ascomycota still remained the absolutely dominant phylum under insect frass addition, while Basidiomycota and Mortierellomycota increased slightly. Over time from day 14 to day 28, Basidiomycota and Others increased slightly under insect frass addition. In the grassland treatment (LN), the 60% field water-holding capacity treatment amended with insect frass showed a high abundance of Ascomycota on day 14, whereas Basidiomycota and Others were relatively higher in the control treatment. On day 28, Ascomycota still maintained a high abundance under insect frass addition. Over time from day 14 to day 28, Ascomycota decreased slightly, whereas Basidiomycota and Others increased slightly under insect frass addition. In the forest land treatment (FT), the 60% field water-holding capacity treatment amended with insect frass showed that the abundance of Ascomycota was higher than that in the control on day 14, whereas the abundance of Basidiomycota was lower than that in the control. On day 28, this change became more obvious: under insect frass addition, Ascomycota further increased, whereas Basidiomycota further decreased; in the control treatment, Basidiomycota consistently maintained a high abundance. Over time from day 14 to day 28, Ascomycota showed an increasing trend, whereas Basidiomycota showed a decreasing trend under insect frass addition.\u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003cp\u003eThe composition and relative abundance of bacterial communities at the genus level under different treatments are shown in Fig.\u0026nbsp;\u003cspan refid=\"Fig4\" class=\"InternalRef\"\u003e4\u003c/span\u003e. In the cropland treatment, under 60% field water-holding capacity (DL 6), the insect frass-amended treatment showed higher abundances of Chryseolinea, Steroidobacter, Pontibacter, Ohtaekwangia, Lysobacter, Nonomuraea, and Chitinophaga than the blank control. From day 14 to day 28, the abundances of Chryseolinea and Steroidobacter further increased in the insect frass-amended treatment, whereas the abundance of Pontibacter decreased slightly, with no significant changes in the blank control. Under 80% field water-holding capacity (DL 8), the insect frass-amended treatment showed increased abundances of Ohtaekwangia and Lysobacter, whereas Ligilactobacillus and Romboutsia decreased; over time, Pontibacter and Ohtaekwangia showed a decreasing trend in the insect frass-amended treatment. In the grassland treatment (LN), under 60% field water-holding capacity, the insect frass-amended treatment showed higher abundances of Lysobacter, Nonomuraea, Chitinophaga, and Steroidobacter than the blank control, whereas Micromonospora and Ligilactobacillus decreased; with prolonged incubation, the abundances of Lysobacter and Micromonospora decreased markedly, whereas those of Nonomuraea and Chitinophaga increased明显. In the forest land treatment (FT), under 60% field water-holding capacity, the insect frass-amended treatment showed markedly higher abundances of Chitinophaga, Sphingomonas, and Burkholderia-Caballeronia-Paraburkholderia than the blank control, whereas Acidothermus, Candidatus Koribacter, Bryobacter, and Candidatus Solibacter decreased. Over time, the abundances of Burkholderia-Caballeronia-Paraburkholderia and Massilia decreased markedly in the insect frass-amended treatment, whereas the blank control showed little change.\u003c/p\u003e \u003cp\u003eThe composition and relative abundance of fungal communities at the genus level under different treatments are shown in Fig.\u0026nbsp;\u003cspan refid=\"Fig5\" class=\"InternalRef\"\u003e5\u003c/span\u003e. In the cropland treatment, under 60% field water-holding capacity (DL 6), the insect frass-amended treatment showed increased abundances of Mortierella and Chaetomiaceae_gen_Incertae_sedis, whereas the abundance of Stachybotrys decreased. From day 14 to day 28, the abundance of Mortierella continued to increase in the insect frass-amended treatment, whereas Stachybotrys in the control treatment increased slightly. In the high-moisture treatment (DL 8), the insect frass-amended treatment showed an increased abundance of Aspergillus, whereas the abundances of Stachybotrys and Chaetomium decreased; over time, the abundances of Zopfiella and Pezizales_gen_Incertae_sedis increased in the insect frass-amended treatment. In the grassland treatment (LN), under 60% field water-holding capacity, insect frass addition markedly increased the abundances of Striaticonidium, Pseudorhypophila, and Humicola, whereas Scedosporium, Gliocephalotrichum, and Oliveonia decreased. With prolonged incubation, Striaticonidium and Humicola decreased slightly, whereas Periconia and Exserohilum increased in the insect frass-amended treatment; in the control treatment, Trichoderma, Fusarium, Scedosporium, and Gliocephalotrichum showed increasing trends, whereas Exserohilum, Neopestalotiopsis, Serendipitaceae_gen_Incertae_sedis, and Ovatospora decreased. In the forest land treatment (FT), under 60% field water-holding capacity, insect frass addition increased the abundance of Penicillium, whereas Trichoderma, Mariannaea, and Saitozyma decreased. Over time, on day 28 compared with day 14, Parachaetomium further increased in the insect frass-amended treatment, whereas the abundances of Trichoderma and Mariannaea decreased in the control treatment.\u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec14\" class=\"Section2\"\u003e \u003ch2\u003e3.4 Correlation Analysis Between Soil Microbial Communities and Physicochemical Factors\u003c/h2\u003e \u003cp\u003eThe correlations among physicochemical indicators and different variable factors, as well as their correlations with bacterial community abundance changes, are shown in Fig.\u0026nbsp;\u003cspan refid=\"Fig6\" class=\"InternalRef\"\u003e6\u003c/span\u003e. The addition of insect frass generally increased the abundances of three bacterial phyla, Pseudomonadota, Bacteroidota, and Chloroflexota, in soil; high soil moisture promoted an increase in the abundance of Chloroflexota, and the abundance of Chloroflexota in soil also increased over time.\u003c/p\u003e \u003cp\u003eIn all treatments, total soluble carbon (TSC) and total soluble organic carbon (TSOC) were significantly positively correlated with the abundance of Actinomycetota; soil ammonium nitrogen (Ammonium) was significantly positively correlated with the abundance of Pseudomonadota; soil nitrate nitrogen (Nitrate) was positively correlated with the abundance of Thermoproteota; soil available potassium (AvbK) was significantly positively correlated with the abundances of Acidobacteriota and Actinomycetota; soil alkali-hydrolyzable nitrogen (AvbN) was significantly positively correlated with the abundances of Pseudomonadota and Actinomycetota; soil available phosphorus (AvbP) was significantly positively correlated with the abundances of Bacteroidota and Gemmatimonadota; soil total nitrogen (TN) was significantly positively correlated with the abundances of Actinomycetota and Bacteroidota; soil total carbon (TC) was significantly positively correlated with the abundances of Actinomycetota and Chloroflexota; and soil pH was significantly positively correlated with the abundances of Acidobacteriota, Gemmatimonadota, Candidatus Eremiobacterota, and Verrucomicrobiota.\u003c/p\u003e \u003cp\u003eThe correlations among physicochemical indicators and different variable factors, as well as their correlations with fungal community abundance changes, are shown in Fig.\u0026nbsp;\u003cspan refid=\"Fig7\" class=\"InternalRef\"\u003e7\u003c/span\u003e. The addition of insect frass generally significantly increased the abundance of Fungi_phy_Incertae_sedis in soil; high soil moisture promoted increases in the abundances of Mucoromycota and Fungi_phy_Incertae_sedis, and the abundance of Fungi_phy_Incertae_sedis in soil also increased over time.\u003c/p\u003e \u003cp\u003eIn all treatments, total soluble carbon (TSC) was significantly positively correlated with the abundance of Glomeromycota; total soluble organic carbon (TSOC) was significantly positively correlated with the abundance of Basidiomycota; soil nitrate nitrogen (Nitrate) was positively correlated with the abundances of Mortierellomycota and Zoopagomycota; soil available potassium (AvbK) was significantly positively correlated with the abundances of Ascomycota, Mortierellomycota, and Mucoromycota; soil alkali-hydrolyzable nitrogen (AvbN) was significantly positively correlated with the abundance of Basidiomycota; soil available phosphorus (AvbP) was significantly positively correlated with the abundances of Ascomycota, Basidiomycota, and Mortierellomycota; soil total nitrogen (TN) and soil total carbon (TC) were significantly positively correlated with the abundance of Glomeromycota; and soil pH was significantly positively correlated with the abundances of Ascomycota, Basidiomycota, and Mucoromycota.\u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec15\" class=\"Section2\"\u003e \u003ch2\u003e3.5 Overall effects on soil after applying mealworm castings\u003c/h2\u003e \u003cp\u003eFigure \u003cspan refid=\"Fig8\" class=\"InternalRef\"\u003e8\u003c/span\u003e shows the PLS-SEM results for the effects of insect frass application on soil properties across the three land-use types under the condition that soil moisture was maintained at 60% of field water-holding capacity. The addition of insect frass reduced total nitrogen and total carbon contents and soluble soil nutrients (soluble carbon and nitrogen) to a certain extent, significantly increased available soil nutrients (available nitrogen, phosphorus, and potassium), and gradually increased soil pH. After the application of mealworm frass, the dominant bacterial phyla in the soil mainly belonged to the functional groups involved in carbon cycling and in nitrogen and phosphorus cycling. Total nutrients, soluble nutrients, and available nutrients were all positively correlated with both microbial functional groups. Among them, total nutrients showed the strongest significant positive correlation with the bacterial communities involved in carbon cycling (\u0026ldquo;Actinomycetota\u0026rdquo;, \u0026ldquo;Bacteroidota\u0026rdquo;, \u0026ldquo;Gemmatimonadota\u0026rdquo;), while soluble nutrients showed the strongest significant positive correlation with the carbon-cycling bacteria (\u0026ldquo;Gemmatimonadota\u0026rdquo;, \u0026ldquo;Pseudomonadota\u0026rdquo;).\u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec16\" class=\"Section2\"\u003e \u003ch2\u003e3.6 The Effect of Mealworm Castings on Soil as Determined by Soil Mechanical Composition\u003c/h2\u003e \u003cp\u003eBy calculating the rate of change in various indicators between the treatment applying mealworm castings and its blank control, SEM was constructed. Soil texture effects on the performance of yellow mealworm castings are shown in Fig.\u0026nbsp;\u003cspan refid=\"Fig9\" class=\"InternalRef\"\u003e9\u003c/span\u003e. Total soil nitrogen and total soil carbon were positively correlated only with sand content, and the correlations were weak. Sand content was significantly positively correlated with soil available nutrients and soil pH (p\u0026thinsp;\u0026lt;\u0026thinsp;0.05), and negatively correlated with total soluble carbon, although the correlation with total soluble carbon was weak. Silt content was significantly positively correlated with soil pH (p\u0026thinsp;\u0026lt;\u0026thinsp;0.05) and negatively correlated with soil available nutrients (p\u0026thinsp;\u0026lt;\u0026thinsp;0.05) and total soluble carbon. Clay content was significantly negatively correlated with soil pH (p\u0026thinsp;\u0026lt;\u0026thinsp;0.05) and positively correlated with total soluble carbon.\u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003c/div\u003e"},{"header":"4 Discussion","content":"\u003cdiv id=\"Sec18\" class=\"Section2\"\u003e \u003ch2\u003e4.1 Effects of Different Treatments on Soil Physicochemical Properties\u003c/h2\u003e \u003cp\u003eAfter mealworm castings were added, changes in soil physicochemical properties differed among land-use types. This variation was partly attributable to differences in baseline soil properties, which caused mealworm castings to affect different soils to different extents. Following the addition of mealworm castings, soil pH generally shifted toward neutrality (approximately 7.0). As an organic amendment, mealworm castings can buffer soil pH and, to some extent, increase pH in acidic soils (Li, D. R. \u003cem\u003eet al\u003c/em\u003e., 2013; Li, Q. J. \u003cem\u003eet al\u003c/em\u003e., 2014). Incorporation of mealworm castings can influence soil pH and nutrient status through multiple pathways. Specifically, mealworm castings can increase soil pH (particularly in acidic soils), enhance soil organic carbon and total nitrogen, and increase available phosphorus and available potassium, while also improving soil structure and microbial activity (Bai, Z. et al., \u003cspan citationid=\"CR14\" class=\"CitationRef\"\u003e2018\u003c/span\u003e; Antoniadis, V. et al., \u003cspan citationid=\"CR15\" class=\"CitationRef\"\u003e2023\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eIn cropland treatments, changes in soil soluble carbon and soluble organic carbon after mealworm casting application showed distinct temporal patterns relative to the blank control under 80% versus 60% of field capacity. Higher moisture can partly slow microbial decomposition of organic matter (Song, D. et al., \u003cspan citationid=\"CR16\" class=\"CitationRef\"\u003e2022\u003c/span\u003e). It also promotes the accumulation of soil ammonium nitrogen while reducing nitrate nitrogen content.\u003c/p\u003e \u003cp\u003eAs an organic soil conditioner, mealworm castings significantly enhance soil easily decomposable carbon content by day 14. Compared to day 14, easily decomposable carbon decreases by day 28, while readily available nutrients begin to increase gradually. This suggests that, during the initial phase after application, mealworm castings undergo preliminary microbial decomposition. Over time, microbial activity progressively mineralizes the castings and releases plant-available nutrients (Zunzunegui, I. et al., \u003cspan citationid=\"CR17\" class=\"CitationRef\"\u003e2024\u003c/span\u003e; Antoniadis, V. et al., \u003cspan citationid=\"CR15\" class=\"CitationRef\"\u003e2023\u003c/span\u003e). However, high moisture content (80% of field capacity) reduced the overall effects of mealworm castings on soil physicochemical properties and slowed microbial mineralization and nutrient release.\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec19\" class=\"Section2\"\u003e \u003ch2\u003e4.2 Effects of Different Treatments on Microbial Community Characteristics\u003c/h2\u003e \u003cp\u003eAfter insect feces was added, the Shannon and Simpson indices increased in forest land (FT) but decreased in croplands (DL) and grassland (LN). These contrasting responses may be driven by multiple factors, including differences in baseline soil conditions, nutrient status, microbial community structure, and niche competition among land-use types (Demenois, J. et al., \u003cspan citationid=\"CR19\" class=\"CitationRef\"\u003e2020\u003c/span\u003e; Huang, Q. et al., \u003cspan citationid=\"CR20\" class=\"CitationRef\"\u003e2024\u003c/span\u003e). In forest land ecosystems, baseline microbial diversity may be relatively low, potentially due to environmental constraints specific to forest ecosystems. When mealworm castings are applied, they act as an organic amendment and supply nutrients such as organic matter, nitrogen, and phosphorus (Li, N. et al., \u003cspan citationid=\"CR21\" class=\"CitationRef\"\u003e2024\u003c/span\u003e; Gurmessa, B. et al., \u003cspan citationid=\"CR22\" class=\"CitationRef\"\u003e2021\u003c/span\u003e). These nutrient inputs may stimulate the growth of microbial taxa that were previously inactive or rare in forest soils, thereby increasing species richness and evenness and ultimately elevating Shannon and Simpson indices (Demenois, J. et al., \u003cspan citationid=\"CR19\" class=\"CitationRef\"\u003e2020\u003c/span\u003e; Li, N. et al., \u003cspan citationid=\"CR21\" class=\"CitationRef\"\u003e2024\u003c/span\u003e). In croplands and grasslands, where baseline diversity is relatively high, nutrients in mealworm castings may preferentially promote the proliferation of already dominant taxa. Moreover, the combined effects of amendment inputs, disturbance to the original soil environment, and potential nutrient surpluses may intensify competitive exclusion, leading to reduced community evenness and, consequently, decreases in the Shannon and Simpson indices (Liu, Z. et al., \u003cspan citationid=\"CR23\" class=\"CitationRef\"\u003e2022\u003c/span\u003e; Wang, X. et al., \u003cspan citationid=\"CR26\" class=\"CitationRef\"\u003e2023\u003c/span\u003e; Zhang, M. et al., \u003cspan citationid=\"CR27\" class=\"CitationRef\"\u003e2024\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eChanges in soil physicochemical properties can strongly shape bacterial community composition. After mealworm feces addition, soil pH, available phosphorus, readily available potassium, and soluble carbon and nitrogen exerted the strongest effects on bacterial communities. Compared with the blank controls, bacterial communities in grassland and forest soils diverged more markedly by day 14, indicating a stronger early-phase response following amendment application. In contrast, bacterial community shifts in cropland soils were less pronounced than those in grassland and forest soils. This difference may be attributable to the relatively higher baseline nutrient status of cropland soils, such that amendment-induced changes in physicochemical properties exert a smaller incremental effect on microbial communities than in forest soils. For example, studies in temperate grassland ecosystems have shown that soil bacterial community composition can be more strongly influenced by nutrient availability than by pH (Zhang, H. et al., \u003cspan citationid=\"CR28\" class=\"CitationRef\"\u003e2024\u003c/span\u003e; Tian, H. et al., \u003cspan citationid=\"CR29\" class=\"CitationRef\"\u003e2017\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eCompared to the control, fungal communities in both forest and grassland soils under the insect dung treatment were more strongly associated with soil available phosphorus, available potassium, and pH on day 14. However, by day 28, the primary factors driving differentiation between the two treatments shifted to soluble carbon, soluble organic carbon, total carbon, ammonium nitrogen and alkaline-hydrolyzable nitrogen. This shift may be explained by temporal changes in substrate availability following amendment addition. Immediately after application, mealworm castings supply abundant nutrients that favor rapid microbial growth. Over time, the introduced carbon is transformed into more labile fractions and undergoes continued mineralization. Consequently, fungal taxa that initially dominate the decomposition of readily available substrates may decline in relative dominance. The functional emphasis of the fungal community may therefore shift from exploiting readily available nutrients toward utilizing more complex carbon sources and/or adapting to conditions with a lower carbon-to-nitrogen ratio (Mao, X. et al., \u003cspan citationid=\"CR30\" class=\"CitationRef\"\u003e2024\u003c/span\u003e).\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec20\" class=\"Section2\"\u003e \u003ch2\u003e4.3 Effects of Different Treatments on Microbial Community Composition\u003c/h2\u003e \u003cp\u003eThe addition of insect feces overall increased the abundance of Pseudomonadota, Actinomycetota, Acidobacteriota, Bacteroidota, Bacillota and Gemmatimonadota within the bacterial community. Mealworm frass is rich in incompletely digested plant polysaccharides, chitin, short-chain fatty acids, and microbial metabolites, which provide soil bacteria with diverse carbon and nitrogen sources (Verardi, A. et al., \u003cspan citationid=\"CR31\" class=\"CitationRef\"\u003e2025\u003c/span\u003e; Nurfikari, A. et al., \u003cspan citationid=\"CR32\" class=\"CitationRef\"\u003e2024\u003c/span\u003e). For example, \u003cem\u003eBacteroidota\u003c/em\u003e and \u003cem\u003ePseudomonadota\u003c/em\u003e, which are capable of degrading complex polysaccharides, as well as \u003cem\u003eActinomycetota\u003c/em\u003e, which can utilize recalcitrant organic matter, may gain a competitive advantage because of the presence of these substrates, thereby increasing in abundance (Nurfikari, A. et al., \u003cspan citationid=\"CR32\" class=\"CitationRef\"\u003e2024\u003c/span\u003e). The addition of mealworm frass alters the physicochemical properties of soil, such as pH, moisture content, redox potential, and oxygen diffusion rate (Nurfikari, A. et al., \u003cspan citationid=\"CR32\" class=\"CitationRef\"\u003e2024\u003c/span\u003e; Nogalska, A. et al., \u003cspan citationid=\"CR33\" class=\"CitationRef\"\u003e2024\u003c/span\u003e). These changes in the microenvironment have selective effects on the growth of different bacterial groups. Mealworm frass may contain bioactive substances produced by the insects themselves, such as antimicrobial peptides, chitinases, and metabolites, and these substances may exert direct or indirect effects on the soil microbial community, for example, by inhibiting certain pathogens or promoting the growth of beneficial bacteria (Rizou, E. et al., \u003cspan citationid=\"CR34\" class=\"CitationRef\"\u003e2025\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eThe addition of mealworm frass increases the abundance of \u003cem\u003eAscomycota\u003c/em\u003e in soil. Mealworm frass contains chitin derived from mealworm molting, which is a nitrogen-containing polysaccharide (Verardi, A. et al., \u003cspan citationid=\"CR31\" class=\"CitationRef\"\u003e2025\u003c/span\u003e; Blakstad, J. I. \u003cem\u003eet al\u003c/em\u003e., 2023). Many groups within \u003cem\u003eAscomycota\u003c/em\u003e, such as \u003cem\u003eFusarium\u003c/em\u003e and \u003cem\u003eTrichoderma\u003c/em\u003e, are known for their strong chitin-degrading ability (Zhang, J. et al., \u003cspan citationid=\"CR36\" class=\"CitationRef\"\u003e2024\u003c/span\u003e). The ability to utilize chitin efficiently may confer a competitive advantage on these ascomycetous fungi in environments amended with mealworm frass, thereby increasing their relative abundance.\u003c/p\u003e \u003cp\u003eOverall, the addition of mealworm frass increased the abundance of \u003cem\u003eChitinophaga\u003c/em\u003e in soil. This phenomenon was mainly attributed to the high chitin content in mealworm frass, which provided sufficient carbon and nitrogen sources for the growth of \u003cem\u003eChitinophaga\u003c/em\u003e (Verardi, A. et al., \u003cspan citationid=\"CR31\" class=\"CitationRef\"\u003e2025\u003c/span\u003e; Zunzunegui, I. et al., \u003cspan citationid=\"CR17\" class=\"CitationRef\"\u003e2024\u003c/span\u003e). In the cropland treatment, insect frass addition promoted increases in the abundances of bacterial communities such as \u003cem\u003eChryseolinea\u003c/em\u003e, \u003cem\u003eSteroidobacter\u003c/em\u003e, \u003cem\u003ePontibacter\u003c/em\u003e, \u003cem\u003eOhtaekwangia\u003c/em\u003e, \u003cem\u003eLysobacter\u003c/em\u003e, \u003cem\u003eNonomuraea\u003c/em\u003e, and \u003cem\u003eChitinophaga\u003c/em\u003e. The increases in these bacterial groups suggest that the soil community was shifting toward enhanced nutrient transformation, biological control, and organic matter mineralization, thereby improving overall soil productivity (Nie, J. et al., \u003cspan citationid=\"CR38\" class=\"CitationRef\"\u003e2025\u003c/span\u003e; Nurfikari, A. et al., \u003cspan citationid=\"CR32\" class=\"CitationRef\"\u003e2024\u003c/span\u003e). In the grassland treatment, insect frass application led to increased abundances of the three saprophytic filamentous fungi \u003cem\u003eStriaticonidium\u003c/em\u003e, \u003cem\u003ePseudorhypophila\u003c/em\u003e, and \u003cem\u003eHumicola\u003c/em\u003e. These fungi play important roles in the decomposition of recalcitrant organic matter (such as lignin and keratin) and in humification, thereby contributing to the accumulation and stabilization of soil organic carbon (Jeffery, S. et al., \u003cspan citationid=\"CR39\" class=\"CitationRef\"\u003e2022\u003c/span\u003e; Hannula, S. E. \u003cem\u003eet al\u003c/em\u003e., 2021). At the same time, the abundances of genera such as \u003cem\u003eScedosporium\u003c/em\u003e, \u003cem\u003eGliocephalotrichum\u003c/em\u003e, and \u003cem\u003eOliveonia\u003c/em\u003e decreased, some of which are considered opportunistic pathogens or weak plant pathogens, suggesting enhanced niche competition and the formation of a disease-suppressive environment (Zhou, W. \u003cem\u003eet al\u003c/em\u003e., 2023; Zhao, W. \u003cem\u003eet al\u003c/em\u003e., 2023).\u003c/p\u003e \u003cp\u003eIn the cropland and grassland treatments, the addition of mealworm frass significantly increased the abundances of \u003cem\u003eStriaticonidium\u003c/em\u003e, \u003cem\u003ePseudorhypophila\u003c/em\u003e, and \u003cem\u003eHumicola\u003c/em\u003e, whereas the abundances of \u003cem\u003eScedosporium\u003c/em\u003e, \u003cem\u003eGliocephalotrichum\u003c/em\u003e, and \u003cem\u003eOliveonia\u003c/em\u003e decreased. In the forest land treatment, the addition of mealworm frass increased the abundance of \u003cem\u003ePenicillium\u003c/em\u003e, whereas the abundances of \u003cem\u003eTrichoderma\u003c/em\u003e, \u003cem\u003eMariannaea\u003c/em\u003e, and \u003cem\u003eSaitozyma\u003c/em\u003e decreased. Increases in fungi such as \u003cem\u003eStriaticonidium\u003c/em\u003e and \u003cem\u003eHumicola\u003c/em\u003e indicate that the degradation and mineralization of soil organic matter were enhanced, which helps accelerate nutrient cycling and convert unstable organic carbon into stable humus, thereby improving soil fertility and carbon sequestration capacity (Yan, H. \u003cem\u003eet al\u003c/em\u003e., 2023; Miao, Y. \u003cem\u003eet al\u003c/em\u003e., 2022). Extracellular enzymes and microbial products secreted by fungi such as Humicola help form and stabilize soil aggregates, improve soil physical structure, enhance soil water- and nutrient-holding capacity, and increase resistance to erosion (Readyhough, T. \u003cem\u003eet al\u003c/em\u003e., 2021; Schroeder, J. \u003cem\u003eet al\u003c/em\u003e., 2021). Increases in beneficial fungi such as Pseudorhypophila and Penicillium, together with decreases in potentially harmful fungi such as Scedosporium, Gliocephalotrichum, and Oliveonia, may enhance soil resistance to pathogens and reduce the risk of plant disease occurrence (Yan, W. et al., \u003cspan citationid=\"CR40\" class=\"CitationRef\"\u003e2024\u003c/span\u003e; Tao, C. \u003cem\u003eet al\u003c/em\u003e., 2023). These changes indicate that mealworm frass, as an organic amendment, can effectively regulate soil microbial communities, particularly fungal communities, thereby optimizing soil ecological functions and supporting sustainable agricultural practices.\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec21\" class=\"Section2\"\u003e \u003ch2\u003e4.4 Effects of Applying Mealworm Castings on Soil\u003c/h2\u003e \u003cp\u003eThrough iterative modeling, we found that the application of mealworm castings introduced abundant nutrients and bioactive substances into the soil, thereby influencing the abundance and activity of the soil microbial community. Microorganisms promoted soil nutrient cycling through their metabolic activities. Following application, microorganisms associated with carbon, nitrogen, and phosphorus cycling became dominant in the soil. These microbes mineralized large organic molecules, converting them into soluble carbon and nitrogen and increasing available phosphorus. Previous studies indicate that mealworm castings effectively elevate available phosphorus levels in soil. Therefore, applying mealworm castings enhances soil nutrient cycling capacity, converting nutrients difficult for plants to utilize into forms \u003cb\u003ethat can be\u003c/b\u003e readily absorbed by plants (Watson, C. et al., \u003cspan citationid=\"CR51\" class=\"CitationRef\"\u003e2021\u003c/span\u003e; Rumbos, C. I. \u003cem\u003eet al\u003c/em\u003e., 2025; Verardi, A. et al., \u003cspan citationid=\"CR31\" class=\"CitationRef\"\u003e2025\u003c/span\u003e). It can also gradually improve soil pH.\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec22\" class=\"Section2\"\u003e \u003ch2\u003e4.5 Effects of Soil Moisture and Mechanical Composition on the Performance of Yellow Mealworm Castings\u003c/h2\u003e \u003cp\u003eBy calculating the relative change rates between the mealworm casting treatment and the blank control and applying iterative modeling, we found that excessively high soil moisture accelerated the increase in soil ammonium nitrogen following mealworm casting application. At the same time, high moisture inhibited the mineralization of mealworm castings by day 14. Only by day 28 did the castings begin to mineralize gradually, at which point soil available nutrients started to increase; however, the increase in soil nitrate nitrogen remained suppressed. Excessive moisture likely promotes ammonification of organic nitrogen by reducing oxygen availability, leading to ammonium accumulation. It may also inhibit aerobic mineralization of organic matter, thereby delaying decomposition. Finally, high moisture strongly suppresses nitrification, limiting nitrate formation (Keiluweit, M. et al., \u003cspan citationid=\"CR44\" class=\"CitationRef\"\u003e2018\u003c/span\u003e; Sun, S. et al., \u003cspan citationid=\"CR45\" class=\"CitationRef\"\u003e2020\u003c/span\u003e; Sriraj, P. et al., \u003cspan citationid=\"CR46\" class=\"CitationRef\"\u003e2022\u003c/span\u003e). Over time, microbial communities may adapt and/or initiate partial anaerobic mineralization, which could restore overall decomposition to some extent; nevertheless, nitrate accumulation is likely to remain constrained under persistently high moisture conditions.\u003c/p\u003e \u003cp\u003eIn soils with different mechanical composition, higher sand contents tend to promote the mineralization of mealworm castings into plant-available nutrients. By contrast, higher silt content was unfavorable for the accumulation of soluble nutrient pools (e.g., total soluble carbon and total soluble nitrogen) and available nutrients. Excessively high clay contents can inhibit both the mineralization of mealworm castings and their pH-modifying effects. Overall, soil texture regulates the mineralization rate of mealworm castings and their effects on soil nutrients and pH by shaping porosity, aeration, water-holding capacity, microbial accessibility to substrates, and pH buffering capacity. Soils with higher sand contents typically have greater porosity and better aeration, which enhance oxygen diffusion and support aerobic microbial activity. In sandy soils, aerobic microorganisms can more efficiently decompose organic matter in mealworm castings and mineralize it into plant-available nutrients such as alkaline-hydrolyzable nitrogen and readily available phosphorus (Ahmed, W. et al., \u003cspan citationid=\"CR47\" class=\"CitationRef\"\u003e2024\u003c/span\u003e; Grzyb, A. et al., \u003cspan citationid=\"CR48\" class=\"CitationRef\"\u003e2020\u003c/span\u003e). Soils with higher silt contents, owing to their relatively large specific surface area, often exhibit stronger adsorption capacity and provide attachment sites for microorganisms, thereby promoting the mineralization of total nitrogen and total carbon in soil (Mao, H.-R. \u003cem\u003eet al\u003c/em\u003e., 2024). In contrast, soils with excessively high clay contents may suppress microbial activity because of poorer aeration and higher water-holding capacity, thereby reducing the decomposition of mealworm castings (Ahmed, W. et al., \u003cspan citationid=\"CR47\" class=\"CitationRef\"\u003e2024\u003c/span\u003e). Moreover, negative charges on clay minerals and organic matter surfaces can adsorb cations, influencing soil buffering capacity and pH dynamics (Jeon, I. et al., \u003cspan citationid=\"CR50\" class=\"CitationRef\"\u003e2023\u003c/span\u003e). In soils with high clay content, the inherently stronger buffering capacity may make pH changes induced by mealworm castings less pronounced. For soils of different textures, their specific characteristics should be considered to optimize the application strategy of mealworm castings, maximizing its ecological benefits as an organic fertilizer source.\u003c/p\u003e \u003c/div\u003e"},{"header":"5 Conclusion","content":"\u003cp\u003eOverall, applying mealworm castings increased various available nutrients in the soil. However, across different land use types, baseline soil nutrient levels affected the effectiveness of mealworm castings. The application of mealworm castings involved relatively low human intervention and showed the most pronounced improvements in relatively nutrient-poor soils, while also producing beneficial effects in nutrient-rich soils. After insect dung application, the abundance of soil saprotrophic microorganisms and other beneficial microbial communities increased to some extent, whereas the abundance of pathogenic microorganisms decreased, which substantially enhanced soil health and improved crop disease resistance. The recommended moisture condition for applying mealworm castings was 60% of field capacity, because excessively high moisture slowed mineralization and decomposition of Tenebrio molitor castings and delayed the time required for them to become effective in soil. Higher sand and silt contents facilitated the mineralization and decomposition of Tenebrio molitor castings, whereas high clay content inhibited the effectiveness of mealworm castings.\u003c/p\u003e \u003cp\u003eThis study elucidated how mealworm castings influenced soil physicochemical properties and microbial communities across soils differing in land-use type, texture, and moisture regime. However, the present work focused on soil responses and did not evaluate outcomes during crop cultivation. Evidence remains limited regarding optimal application rates required to maximize overall benefits under different land-use types and moisture conditions. In addition, the potential role of leaching was not considered when assessing texture-dependent effects. Future studies should address these gaps through complementary experiments to provide more robust and comprehensive guidance for the agricultural use of mealworm castings.\u003c/p\u003e "},{"header":"Declarations","content":"\u003cp\u003e \u003cb\u003eAuthor Responsibility Statement\u003c/b\u003e \u003c/p\u003e \u003cp\u003eWe, the authors of this paper, hereby solemnly declare once more that we confirm this research constitutes original work, with all experimental data being first-hand data obtained through standardised experiments. This study has not involved any form of academic misconduct, including but not limited to data falsification, plagiarism, multiple submissions, or duplicate publication. All references have been properly cited.\u003c/p\u003e\u003cp\u003e \u003ch2\u003eConflict of Interest Statement\u003c/h2\u003e \u003cp\u003eWe declare that there are no competing financial interests or personal relationships relevant to this research.\u003c/p\u003e \u003c/p\u003e\u003ch2\u003eAuthor Contribution\u003c/h2\u003e\u003cp\u003eZ:Responsible for experimental design, sample collection and mechanical composition analysis; completed moisture infiltration rate measurements and data collation for the earthworm compost treatment group; drafted the initial manuscript. Z:Participated in determining soil physical and chemical properties, sample collection, assisted in collating data across different land use types, conducted data plotting, chart visualisation, and paper revisions. L:Responsible for earthworm biomass monitoring and soil microbial community analysis, collated comparative data across different land use types and performed data processing. W and L:Assisted with earthworm biomass monitoring and preliminary microbial community analysis. J and J:Assisted with soil sample collection and data organisation. Z:As supervising tutor, provided research direction, land use typedata support, and field sampling technical guidance. X:As corresponding author, guided research direction, provided conceptual and technical support, and reviewed the manuscript.\u003c/p\u003e\u003ch2\u003eAcknowledgements\u003c/h2\u003e \u003cp\u003eThis work was supported by the Natural Science Foundation of Guangdong Province (2023A1515010593).\u003c/p\u003e\u003ch2\u003eData Availability\u003c/h2\u003e\u003cp\u003eAll data generated or analyzed during this study are included in this published article and its supplementary information files\u003c/p\u003e"},{"header":"References","content":"\u003col\u003e\u003cli\u003e\u003cspan\u003eHe L, Zhang Y, Ding M et al (2021) Sustainable strategy for lignocellulosic crop wastes reduction by \u003cem\u003eTenebrio molitor\u003c/em\u003e linnaeus (mealworm) and potential use of mealworm frass as a fertilizer[J]. 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China Agriculture, Beijing\u003c/span\u003e\u003c/li\u003e\u003c/ol\u003e"}],"fulltextSource":"","fullText":"","funders":[],"hasAdminPriorityOnWorkflow":false,"hasManuscriptDocX":true,"hasOptedInToPreprint":true,"hasPassedJournalQc":"","hasAnyPriority":false,"hideJournal":true,"highlight":"","institution":"","isAcceptedByJournal":false,"isAuthorSuppliedPdf":false,"isDeskRejected":"","isHiddenFromSearch":false,"isInQc":false,"isInWorkflow":false,"isPdf":false,"isPdfUpToDate":true,"isWithdrawnOrRetracted":false,"journal":{"display":true,"email":"[email protected]","identity":"researchsquare","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":true,"externalIdentity":"","sideBox":"","snPcode":"","submissionUrl":"/submission","title":"Research Square","twitterHandle":"researchsquare","acdcEnabled":true,"dfaEnabled":false,"editorialSystem":"","reportingPortfolio":"","inReviewEnabled":false,"inReviewRevisionsEnabled":true},"keywords":"Tenebrio molitor castings, organic fertilizer, soil amendment, soil microbial community structure, amplicon sequencing","lastPublishedDoi":"10.21203/rs.3.rs-8998912/v1","lastPublishedDoiUrl":"https://doi.org/10.21203/rs.3.rs-8998912/v1","license":{"name":"CC BY 4.0","url":"https://creativecommons.org/licenses/by/4.0/"},"manuscriptAbstract":"\u003cp\u003eTo investigate the effects of mealworm casting amendment on different land-use soils and find out the optimal soil moisture content for application, three distinct soils of different soil texture were selected for soil incubation experiment, one of which was set to two moisture levels. Soil samples were collected on days 14 and 28 of the incubation. Research findings indicate that soil mealworm casting amendemnt stimulates soil carbon and nitrogen cycling, facilitating the conversion of nutrients into forms readily absorbed by plants. The most pronounced effect is observed in soil ammonium nitrogen content, with an average increase of 620.7%; the average increases in soluble carbon and soluble organic carbon were 80.7% and 92.5%, respectively. Adding insect manure significantly increased the abundance of microorganisms involved in the soil carbon cycling, such as Proteobacteria and Bacteroidetes. The mealworm manure also increased beneficial taxa such as \u003cem\u003eChitinophaga\u003c/em\u003e and suppressed disease-associated microorganisms, thereby improving the soil microbial community structure. Moisture conditions and soil mechanical composition were key factors influencing the effectiveness of mealworm castings. The optimal moisture content for mealworm castings application is 60% of the field capacity, while higher moisture (80% of the field capacity) inhibits casting mineralization and reduces soil nitrate nitrogen content by 73.2% at day 28. Suitable moisture contents promoted mineralization of \u003cem\u003eTenebrio molitor\u003c/em\u003e castings, whereas excessive clay content constrained both mineralization and the frass-induced increase in soil pH.\u003c/p\u003e","manuscriptTitle":"Soil Mechanical Composition and Moisture Enhanced the Improvement of Mealworm Casting Amendment on Different Land Use Soils","msid":"","msnumber":"","nonDraftVersions":[{"code":1,"date":"2026-04-06 12:32:45","doi":"10.21203/rs.3.rs-8998912/v1","editorialEvents":[{"type":"communityComments","content":0}],"status":"published","journal":{"display":true,"email":"[email protected]","identity":"researchsquare","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":true,"externalIdentity":"","sideBox":"","snPcode":"","submissionUrl":"/submission","title":"Research Square","twitterHandle":"researchsquare","acdcEnabled":true,"dfaEnabled":false,"editorialSystem":"","reportingPortfolio":"","inReviewEnabled":false,"inReviewRevisionsEnabled":true}}],"origin":"","ownerIdentity":"2ffcc410-fcc1-467b-91f2-f642f9b0d7be","owner":[],"postedDate":"April 6th, 2026","published":true,"recentEditorialEvents":[],"rejectedJournal":[],"revision":"","amendment":"","status":"posted","subjectAreas":[],"tags":[],"updatedAt":"2026-04-27T06:11:29+00:00","versionOfRecord":[],"versionCreatedAt":"2026-04-06 12:32:45","video":"","vorDoi":"","vorDoiUrl":"","workflowStages":[]},"version":"v1","identity":"rs-8998912","journalConfig":"researchsquare"},"__N_SSP":true},"page":"/article/[identity]/[[...version]]","query":{"redirect":"/article/rs-8998912","identity":"rs-8998912","version":["v1"]},"buildId":"XKTyCvWXoU3ODBz1xrDgd","isFallback":false,"isExperimentalCompile":false,"dynamicIds":[84888],"gssp":true,"scriptLoader":[]}